Tyrannosaurus - Wikipedia 01/01/2025, 00:07

Tyrannosaurus
(Redirected from Tyrannosaurus rex)

Tyrannosaurus (/tɪˌrænəˈsɔːrəs, taɪ-/)[a] is a genus

of large theropod dinosaur. The type species Tyrannosaurus
Tyrannosaurus rex (rex meaning 'king' in Latin), Temporal range: Late Cretaceous,
often shortened to T. rex or colloquially T-Rex, is one
of the best represented theropods. It lived throughout
what is now western North America, on what was then
an island continent known as Laramidia.
Tyrannosaurus had a much wider range than other
tyrannosaurids. Fossils are found in a variety of rock
formations dating to the latest Campanian-
Maastrichtian ages of the late Cretaceous period, 72.7 to
66 million years ago. It was the last known member of
Reconstruction of the T. rex type
the tyrannosaurids and among the last non-avian
specimen at the Carnegie Museum of
dinosaurs to exist before the Cretaceous–Paleogene
Natural History
extinction event.
Scientific classification
Like other tyrannosaurids, Tyrannosaurus was a
bipedal carnivore with a massive skull balanced by a Domain: Eukaryota
long, heavy tail. Relative to its large and powerful hind Kingdom: Animalia
limbs, the forelimbs of Tyrannosaurus were short but
Phylum: Chordata
unusually powerful for their size, and they had two
clawed digits. The most complete specimen measures Clade: Dinosauria
12.3–12.4 m (40–41 ft) in length, but according to most Clade: Saurischia
modern estimates, Tyrannosaurus could have exceeded
Clade: Theropoda
sizes of 13 m (43 ft) in length, 3.7–4 m (12–13 ft) in hip
height, and 8.8 t (8.7 long tons; 9.7 short tons) in mass. Family: †Tyrannosauridae
Although some other theropods might have rivaled or Subfamily: †Tyrannosaurinae
exceeded Tyrannosaurus in size, it is still among the
largest known land predators, with its estimated bite Clade: †Tyrannosaurini
force being the largest among all terrestrial animals. By Genus: †Tyrannosaurus
far the largest carnivore in its environment, Osborn, 1905
Tyrannosaurus rex was most likely an apex predator,
preying upon hadrosaurs, juvenile armored herbivores Type species
like ceratopsians and ankylosaurs, and possibly †Tyrannosaurus rex
sauropods. Some experts have suggested the dinosaur

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was primarily a scavenger. The question of whether Osborn, 1905

Tyrannosaurus was an apex predator or a pure
Other species
scavenger was among the longest debates in
paleontology. Most paleontologists today accept that †T. mcraeensis Dalman et al.,
Tyrannosaurus was both a predator and a scavenger.
2024
Specimens of Tyrannosaurus rex include some that are See text
nearly complete skeletons. Soft tissue and proteins have
Synonyms
been reported in at least one of these specimens. The
abundance of fossil material has allowed significant Genus synonymy
research into many aspects of its biology, including its Dinotyrannus
life history and biomechanics. The feeding habits,
Olshevsky, 1995
physiology, and potential speed of Tyrannosaurus rex
are a few subjects of debate. Its taxonomy is also Dynamosaurus
controversial, as some scientists consider Tarbosaurus Osborn, 1905
bataar from Asia to be a third Tyrannosaurus species, Manospondylus
while others maintain Tarbosaurus is a separate genus. Cope, 1892
Several other genera of North American tyrannosaurids
have also been synonymized with Tyrannosaurus. At Nanotyrannus?
present, two species of Tyrannosaurus are considered Bakker, Williams & Currie, 1988
valid; the type species, T. rex, and the earlier in age Stygivenator
and more recently discovered T. mcraeensis. Olshevsky, 1995

As the archetypal theropod, Tyrannosaurus has been Species synonymy

one of the best-known dinosaurs since the early 20th Aublysodon amplus?
century and has been featured in film, advertising, Marsh, 1892
postal stamps, and many other media.
Deinodon amplus?
(Marsh, 1892) Hay, 1902

History of research Manospondylus amplus?

(Marsh, 1892) Olshevsky, 1978
Stygivenator amplus?
Earliest finds (Marsh, 1892) Olshevsky, 1995
Tyrannosaurus amplus?
(Marsh, 1892) Hay, 1930
Aublysodon cristatus?
Marsh, 1892
Deinodon cristatus?
(Marsh, 1892) Hay, 1902
Type specimen (AMNH 3982) of
Stygivenator cristatus?
Manospondylus gigas
(Marsh, 1892) Olshevsky, 1995

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A tooth from what is now documented as a Ornithomimus grandis

Tyrannosaurus rex was found in July 1874 upon South Marsh, 1890
Table Mountain (Colorado) by Jarvis Hall (Colorado)
student Peter T. Dotson under the auspices of Prof. Manospondylus gigas
Arthur Lakes near Golden, Colorado.[1] In the early Cope, 1892
1890s, John Bell Hatcher collected postcranial elements Dynamosaurus imperiosus
in eastern Wyoming. The fossils were believed to be Osborn, 1905
from the large species Ornithomimus grandis (now
Tyrannosaurus imperiosus
Deinodon) but are now considered T. rex remains.[2]
(Osborn, 1905) Swinton, 1970
In 1892, Edward Drinker Cope found two vertebral Gorgosaurus lancensis
fragments of a large dinosaur. Cope believed the Gilmore, 1946
fragments belonged to an "agathaumid" (ceratopsid)
Albertosaurus lancensis
dinosaur, and named them Manospondylus gigas,
meaning "giant porous vertebra", in reference to the (Gilmore, 1946) Russell, 1970
numerous openings for blood vessels he found in the Deinodon lancensis
bone.[2] The M. gigas remains were, in 1907, identified (Gilmore, 1946) Kuhn, 1965
by Hatcher as those of a theropod rather than a
Aublysodon lancensis
ceratopsid.[3]
(Gilmore, 1946) Charig in Appleby,
Henry Fairfield Osborn recognized the similarity Charig, Cox, Kermack & Tarlo, 1967
between Manospondylus gigas and T. rex as early as Nanotyrannus lancensis
1917, by which time the second vertebra had been lost. (Gilmore, 1946) Bakker, Williams &
Owing to the fragmentary nature of the Manospondylus
Currie, 1988
vertebrae, Osborn did not synonymize the two genera,
instead considering the older genus indeterminate.[4] Albertosaurus "megagracilis"
In June 2000, the Black Hills Institute found around Paul, 1988a (nomen nudum)
10% of a Tyrannosaurus skeleton (BHI 6248) at a site Dinotyrannus megagracilis
that might have been the original M. gigas locality.[5] Olshevsky, 1995
Aublysodon molnaris
Skeleton discovery and naming Paul, 1988a
Barnum Brown, assistant curator of the American Aublysodon molnari
Museum of Natural History, found the first partial Paul, 1988a emend Paul, 1990
skeleton of T. rex in eastern Wyoming in 1900. Brown
Stygivenator molnari
found another partial skeleton in the Hell Creek
Formation in Montana in 1902, comprising (Paul, 1988a emend Paul, 1990)
approximately 34 fossilized bones.[6] Writing at the Olshevsky, 1995
time Brown said "Quarry No. 1 contains the femur,
pubes, humerus, three vertebrae and two undetermined bones of a large Carnivorous Dinosaur
not described by Marsh. ... I have never seen anything like it from the Cretaceous."[7] Henry
Fairfield Osborn, president of the American Museum of Natural History, named the second
skeleton T. rex in 1905. The generic name is derived from the Greek words τύραννος (tyrannos,

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meaning "tyrant") and σαῦρος (sauros, meaning "lizard").

Osborn used the Latin word rex, meaning "king", for the
specific name. The full binomial therefore translates to
"tyrant lizard the king" or "King Tyrant Lizard",
emphasizing the animal's size and presumed dominance
over other species of the time.[6]

Osborn named the other

specimen
Dynamosaurus
imperiosus in a paper in
1905.[6] In 1906, Osborn Outdated skeletal restoration by
Dynamosaurus imperiosus holotype
(NHMUK PV R8020, previously
recognized that the two William D. Matthew from 1905,

AMNH 5886) on display at the skeletons were from the published alongside Osborn's
description paper
Natural History Museum same species and
selected Tyrannosaurus
as the preferred name.[8] In 1941, the T. rex type specimen
was sold to the Carnegie Museum of Natural History in Pittsburgh, Pennsylvania, for $7,000.[7]
The original Dynamosaurus material now resides in the collections of the Natural History
Museum, London.[9] Dynamosaurus would later be honored by the 2018 description of another
species of tyrannosaurid by Andrew McDonald and colleagues, Dynamoterror dynastes, whose
name was chosen in reference to the 1905 name, as it had been a "childhood favorite" of
McDonald's.[10]

From the 1910s through the end of the 1950s, Barnum's discoveries remained the only
specimens of Tyrannosaurus, as the Great Depression and wars kept many paleontologists out
of the field.[5]

Resurgent interest
Beginning in the 1960s, there was renewed interest in
Tyrannosaurus, resulting in the recovery of 42 skeletons
(5–80% complete by bone count) from Western North
America.[5] In 1967, Dr. William MacMannis located and
recovered the skeleton named "MOR 008", which is 15%
complete by bone count and has a reconstructed skull
displayed at the Museum of the Rockies. The 1990s saw
numerous discoveries, with nearly twice as many finds as in Specimen "Sue", Field Museum of
all previous years, including two of the most complete Natural History, Chicago
skeletons found to date: Sue and Stan.[5]

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Sue Hendrickson, an amateur paleontologist, discovered the most complete (approximately

85%) and largest Tyrannosaurus skeleton in the Hell Creek Formation on August 12, 1990. The
specimen Sue, named after the discoverer, was the object of a legal battle over its ownership. In
1997, the litigation was settled in favor of Maurice Williams, the original land owner. The fossil
collection was purchased by the Field Museum of Natural History at auction for $7.6 million,
making it the most expensive dinosaur skeleton until the sale of Stan for $31.8 million in
2020.[11] From 1998 to 1999, Field Museum of Natural History staff spent over 25,000 hours
taking the rock off the bones.[12] The bones were then shipped to New Jersey where the mount
was constructed, then shipped back to Chicago for the final assembly. The mounted skeleton
opened to the public on May 17, 2000, in the Field Museum of Natural History. A study of this
specimen's fossilized bones showed that Sue reached full size at age 19 and died at the age of 28,
the longest estimated life of any tyrannosaur known.[13]

Another Tyrannosaurus, nicknamed Stan (BHI 3033), in

honor of amateur paleontologist Stan Sacrison, was
recovered from the Hell Creek Formation in 1992. Stan is
the second most complete skeleton found, with 199 bones
recovered representing 70% of the total.[14] This
"Scotty", the largest known tyrannosaur also had many bone pathologies, including
specimen, exhibited in Japan broken and healed ribs, a broken (and healed) neck, and a
substantial hole in the back of its head, about the size of a
Tyrannosaurus tooth.[15]

In 1998, 20-year-old Bucky Derflinger noticed a T. rex toe exposed above ground, making him
the youngest person to discover a Tyrannosaurus. The specimen, dubbed Bucky in honor of its
discoverer, was a young adult, 3.0 metres (10 ft) tall and 11 metres (35 ft) long. Bucky is the first
Tyrannosaurus to be found that preserved a furcula (wishbone). Bucky is permanently
displayed at The Children's Museum of Indianapolis.[16]

In the summer of 2000, crews organized by Jack Horner

discovered five Tyrannosaurus skeletons near the Fort Peck
Reservoir.[17] In 2001, a 50% complete skeleton of a juvenile
Tyrannosaurus was discovered in the Hell Creek Formation
by a crew from the Burpee Museum of Natural History.
The specimens "Sue", AMNH 5027,
Dubbed Jane (BMRP 2002.4.1), the find was thought to be "Stan", and "Jane", to scale with a
the first known skeleton of a pygmy tyrannosaurid, human.
Nanotyrannus, but subsequent research revealed that it is
more likely a juvenile Tyrannosaurus, and the most
complete juvenile example known;[18] Jane is exhibited at the Burpee Museum of Natural
History.[19] In 2002, a skeleton nicknamed "Wyrex", discovered by amateur collectors Dan
Wells and Don Wyrick, had 114 bones and was 38% complete. The dig was concluded over 3
weeks in 2004 by the Black Hills Institute with the first live online Tyrannosaurus excavation
providing daily reports, photos, and video.[5]

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In 2006, Montana State University revealed that it possessed the largest Tyrannosaurus skull
yet discovered (from a specimen named MOR 008), measuring 5 feet (152 cm) long.[20]
Subsequent comparisons indicated that the longest head was 136.5 centimetres (53.7 in) (from
specimen LACM 23844) and the widest head was 90.2 centimetres (35.5 in) (from Sue).[21]

Footprints
Two isolated fossilized footprints have been tentatively
assigned to T. rex. The first was discovered at Philmont
Scout Ranch, New Mexico, in 1983 by American geologist
Charles Pillmore. Originally thought to belong to a
hadrosaurid, examination of the footprint revealed a large
'heel' unknown in ornithopod dinosaur tracks, and traces of
what may have been a hallux, the dewclaw-like fourth digit
of the tyrannosaur foot. The footprint was published as the
Probable footprint from New Mexico ichnogenus Tyrannosauripus pillmorei in 1994, by Martin
Lockley and Adrian Hunt. Lockley and Hunt suggested that
it was very likely the track was made by a T. rex, which
would make it the first known footprint from this species. The track was made in what was once
a vegetated wetland mudflat. It measures 83 centimeters (33 in) long by 71 centimeters (28 in)
wide.[22]

A second footprint that may have been made by a Tyrannosaurus was first reported in 2007 by
British paleontologist Phil Manning, from the Hell Creek Formation of Montana. This second
track measures 72 centimeters (28 in) long, shorter than the track described by Lockley and
Hunt. Whether or not the track was made by Tyrannosaurus is unclear, though Tyrannosaurus
is the only large theropod known to have existed in the Hell Creek Formation.[23][24]

A set of footprints in Glenrock, Wyoming dating to the Maastrichtian stage of the Late
Cretaceous and hailing from the Lance Formation were described by Scott Persons, Phil Currie
and colleagues in 2016, and are believed to belong to either a juvenile T. rex or the dubious
tyrannosaurid Nanotyrannus lancensis. From measurements and based on the positions of the
footprints, the animal was believed to be traveling at a walking speed of around 2.8 to 5 miles
per hour and was estimated to have a hip height of 1.56 to 2.06 m (5.1 to 6.8 ft).[25][26][27] A
follow-up paper appeared in 2017, increasing the speed estimations by 50–80%.[28]

Description

Size

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T. rex was one of the largest land

carnivores of all time. One of its largest
and the most complete specimens,
nicknamed Sue (FMNH PR2081), is
located at the Field Museum of Natural
History in Chicago. Sue measured 12.3–
12.4 m (40–41 ft) long,[29][30] was 3.66–
3.96 m (12.0–13.0 ft) tall at the Size (in blue) compared to select giant theropods and a
human
hips,[31][32][33] and according to the most
recent studies, using a variety of
techniques, maximum body masses have been estimated approximately 8.4–8.46 t (8.27–8.33
long tons; 9.26–9.33 short tons).[34][35] A specimen nicknamed Scotty (RSM P2523.8), located
at the Royal Saskatchewan Museum, is reported to measure 13 m (43 ft) in length. Using a mass
estimation technique that extrapolates from the circumference of the femur, Scotty was
estimated as the largest known specimen at 8.87 t (8.73 long tons; 9.78 short tons) in body
mass.[34][36]

Not every adult Tyrannosaurus specimen recovered is as big. Historically average adult mass
estimates have varied widely over the years, from as low as 4.5 t (4.4 long tons; 5.0 short
tons),[37][38] to more than 7.2 t (7.1 long tons; 7.9 short tons),[39] with most modern estimates
ranging between 5.4 and 8.0 t (5.3 and 7.9 long tons; 6.0 and 8.8 short tons).[29][40][41][42][43]

A 2024 study found that there was little evidence of size-based sexual dimorphism in T. rex.[44]

Skull
The largest known T. rex skulls measure up to 1.54 m (5.1 ft)
in length.[20][31] Large fenestrae (openings) in the skull
reduced weight, as in all carnivorous theropods. In other
respects Tyrannosaurus's skull was significantly different
from those of large non-tyrannosaurid theropods. It was
extremely wide at the rear but had a narrow snout, allowing
unusually good binocular vision.[45][46] The skull bones
were massive and the nasals and some other bones were
Profile view of a skull (AMNH 5027) fused, preventing movement between them; but many were
pneumatized (contained a "honeycomb" of tiny air spaces)
and thus lighter. These and other skull-strengthening
features are part of the tyrannosaurid trend towards an increasingly powerful bite, which easily
surpassed that of all non-tyrannosaurids.[47][48][49] The tip of the upper jaw was U-shaped
(most non-tyrannosauroid carnivores had V-shaped upper jaws), which increased the amount
of tissue and bone a tyrannosaur could rip out with one bite, although it also increased the
stresses on the front teeth.[50]

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The teeth of T. rex displayed marked heterodonty

(differences in shape).[51][52] The premaxillary teeth, four
per side at the front of the upper jaw, were closely packed,
D-shaped in cross-section, had reinforcing ridges on the
rear surface, were incisiform (their tips were chisel-like
blades) and curved backwards. The D-shaped cross-section,
reinforcing ridges and backwards curve reduced the risk
that the teeth would snap when Tyrannosaurus bit and
pulled. The remaining teeth were robust, like "lethal
Skull replica of specimen "Sue",
bananas" rather than daggers, more widely spaced and also showing dentition
had reinforcing ridges.[53] Those in the upper jaw, twelve
per side in mature individuals,[51] were larger than their
counterparts of the lower jaw, except at the rear. The largest found so far is estimated to have
been 30.5 cm (12.0 in) long including the root when the animal was alive, making it the largest
tooth of any carnivorous dinosaur yet found.[54] The lower jaw was robust. Its front dentary
bone bore thirteen teeth. Behind the tooth row, the lower jaw became notably taller.[51] The
upper and lower jaws of Tyrannosaurus, like those of many dinosaurs, possessed numerous
foramina, or small holes in the bone. Various functions have been proposed for these foramina,
such as a crocodile-like sensory system[55] or evidence of extra-oral structures such as scales or
potentially lips,[56][57][58] with subsequent research on theropod tooth wear patterns
supporting such a proposition.[59]

Skeleton
The vertebral column of Tyrannosaurus consisted of ten
neck vertebrae, thirteen back vertebrae and five sacral
vertebrae. The number of tail vertebrae is unknown and
could well have varied between individuals but probably
Life restoration showing scaly skin with numbered at least forty. Sue was mounted with forty-
sparse feathering, and lipped jaws
seven of such caudal vertebrae.[51] The neck of T. rex
formed a natural S-shaped curve like that of other
theropods. Compared to these, it was exceptionally short,
deep and muscular to support the massive head. The
second vertebra, the axis, was especially short. The
remaining neck vertebrae were weakly opisthocoelous,
i.e. with a convex front of the vertebral body and a
Skeletal reconstruction of specimen
"Sue" concave rear. The vertebral bodies had single
pleurocoels, pneumatic depressions created by air sacs,
on their sides.[51] The vertebral bodies of the torso were
robust but with a narrow waist. Their undersides were keeled. The front sides were concave with
a deep vertical trough. They had large pleurocoels. Their neural spines had very rough front and
rear sides for the attachment of strong tendons. The sacral vertebrae were fused to each other,

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both in their vertebral bodies and neural spines. They were pneumatized. They were connected
to the pelvis by transverse processes and sacral ribs. The tail was heavy and moderately long, in
order to balance the massive head and torso and to provide space for massive locomotor
muscles that attached to the thighbones. The thirteenth tail vertebra formed the transition point
between the deep tail base and the middle tail that was stiffened by a rather long front
articulation processes. The underside of the trunk was covered by eighteen or nineteen pairs of
segmented belly ribs.[51]

The shoulder girdle was longer than the entire forelimb. The
shoulder blade had a narrow shaft but was exceptionally
expanded at its upper end. It connected via a long forward
protrusion to the coracoid, which was rounded. Both
shoulder blades were connected by a small furcula. The
paired breast bones possibly were made of cartilage only.[51]

The forelimb or arm was very short. The upper arm bone,
the humerus, was short but robust. It had a narrow upper
end with an exceptionally rounded head. The lower arm
bones, the ulna and radius, were straight elements, much
shorter than the humerus. The second metacarpal was Right forelimb of specimen "Sue"
longer and wider than the first, whereas normally in
theropods the opposite is true. The forelimbs had only two
clawed fingers,[51] along with an additional splint-like small third metacarpal representing the
remnant of a third digit.[60]

The pelvis was a large structure. Its upper bone, the ilium,
was both very long and high, providing an extensive
attachment area for hindlimb muscles. The front pubic bone
ended in an enormous pubic boot, longer than the entire
shaft of the element. The rear ischium was slender and
straight, pointing obliquely to behind and below.[51]

In contrast to the arms, the hindlimbs were among the

longest in proportion to body size of any theropod. In the
Pelvic girdle of specimen MOR 555
foot, the metatarsus was "arctometatarsalian", meaning that
the part of the third metatarsal near the ankle was pinched.
The third metatarsal was also exceptionally sinuous.[51] Compensating for the immense bulk of
the animal, many bones throughout the skeleton were hollowed, reducing its weight without
significant loss of strength.[51]

Classification

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Tyrannosaurus is the type genus of the superfamily

Tyrannosauroidea, the family Tyrannosauridae, and the
subfamily Tyrannosaurinae; in other words it is the
standard by which paleontologists decide whether to include
other species in the same group. Other members of the
tyrannosaurine subfamily include the North American
Daspletosaurus and the Asian Tarbosaurus,[18][61] both of
which have occasionally been synonymized with
Tyrannosaurus.[62]
Skull casts of different
Tyrannosaurids were once commonly thought to be Tyrannosaurus specimens
descendants of earlier large theropods such as megalosaurs
and carnosaurs, although more recently they were
reclassified with the generally smaller coelurosaurs.[50] The earliest tyrannosaur group were the
crested proceratosaurids, while later and more derived members belong to the
Pantyrannosauria. Tyrannosaurs started out as small theropods; however at least some became
larger by the Early Cretaceous.

Tyrannosauroids are characterized by their fused nasals and dental arrangement.

Pantyrannosaurs are characterized by unique features in their hips as well as an enlarged
foramen in the quadrate, a broad postorbital and hourglass shaped nasals. Some of the more
derived pantyrannosaurs lack nasal pneumaticity and have a lower humerus to femur ratio with
their arms starting to see some reduction. Some pantyrannosaurs started developing an
arctometatarsus. Eutyrannosaurs have a rough texture on their nasal bones and their
mandibular fenestra is reduced externally. Tyrannosaurids lack kinetic skulls or special crests
on their nasal bones, and have a lacrimal with a distinctive process on it. Tyrannosaurids also
have an interfenestral strut that is less than half as big as the maxillary fenestra.[63]

It is quite likely that tyrannosauroids rose to prominence after the decline in allosauroid and
megalosauroid diversity seen during the early stages of the Late Cretaceous. Below is a simple
cladogram of general tyrannosauroid relationships that was found after an analysis conducted
by Li and colleagues in 2009.[64]

Tyrannosauroidea
Guanlong

Proceratosaurus

Pantyrannosauria
Dilong

Eotyrannus

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Xiongguanlong

Eutyrannosauria Appalachiosaurus

Tyrannosauridae

Many phylogenetic analyses have found Tarbosaurus bataar to be the sister taxon of T. rex.[61]
The discovery of the tyrannosaurid Lythronax further indicates that Tarbosaurus and
Tyrannosaurus are closely related, forming a clade with fellow Asian tyrannosaurid
Zhuchengtyrannus, with Lythronax being their sister taxon.[65][66] A further study from 2016
by Steve Brusatte, Thomas Carr and colleagues, also indicates that Tyrannosaurus may have
been an immigrant from Asia, as well as a possible descendant of Tarbosaurus.[67]

Below is the cladogram of Tyrannosauridae based on the phylogenetic analysis conducted by

Loewen and colleagues in 2013.[65]

Tyrannosauridae Albertosaurinae
Gorgosaurus libratus

Albertosaurus sarcophagus

Tyrannosaurinae Dinosaur Park tyrannosaurid

Daspletosaurus torosus

Two Medicine tyrannosaurid

Teratophoneus curriei

Bistahieversor sealeyi

Lythronax argestes

Tyrannosaurus rex

Tarbosaurus bataar

Zhuchengtyrannus magnus

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In their 2024 description of Tyrannosaurus mcraeensis, Dalman et al. recovered similar results
to previous analyses, with Tyrannosaurus as the sister taxon to the clade formed by
Tarbosaurus and Zhuchengtyrannus, called the Tyrannosaurini. They also found support for a
monophyletic clade containing Daspletosaurus and Thanatotheristes, typically referred to as
the Daspletosaurini.[68][69]

Albertosaurus sarcophagus

Gorgosaurus libratus

Daspletosaurus horneri

Thanatotheristes

Daspletosaurus torosus

Daspletosaurus wilsoni

Teratophoneus

Nanuqsaurus

Bistahieversor

Lythronax

Tyrannosaurini Tyrannosaurus mcraeensis

Tyrannosaurus rex

Zhuchengtyrannus

Tarbosaurus

Additional species
In 1955, Soviet paleontologist Evgeny Maleev named a new species, Tyrannosaurus bataar,
from Mongolia.[70] By 1965, this species was renamed as a distinct genus, Tarbosaurus
bataar.[71] While most palaeontologists continue to maintain the two as distinct genera, some
authors such as Thomas Holtz, Kenneth Carpenter, and Thomas Carr argue that the two species

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are similar enough to be considered members of the same

genus, restoring the Mongolian taxon's original binomial
name.[50][72][55]

Some specimens from the Late Cretaceous deposits of China

have been described as new species of Tyrannosaurus: T.
lanpingensis based on isolated lateral tooth from the red
beds of Yunnan in 1975; T. turpanensis from the Subashi
Formation, Turpan Basin, Xinjiang in 1978; and T.
luanchuanensis from the Quiba Formation, Tantou Basin,
Henan Province in 1979–1980.[73][74][75] All these taxa were
published without detailed descriptions and were later Diagram showing the differences
accepted as junior synonyms of Tarbosaurus bataar by between a generalized Tarbosaurus
(A) and Tyrannosaurus rex (B) skull
Holtz in 2004.[61][74][75]

VGI, no. 231/3, a large phalanx bone, assigned to Tyrannosaurus sp. by Yarkov in 2000, was
found in the Lower Maastrichtian of Bereslavka, Russia. In 2004, Averianov and Yarkov
reinterpreted it as a metacarpal I or metatarsal I that possibly belongs to ceratosaur.[76] In their
2023 overview, Averianov and Lopatin mention this specimen as well as a single tooth from the
same site only as Theropoda indet.[77]

In 2001, various tyrannosaurid teeth and a metatarsal unearthed in a quarry near Zhucheng,
China were assigned by Chinese paleontologist Hu Chengzhi to the newly erected species
Tyrannosaurus zhuchengensis. However, in a nearby site, a right maxilla and left jawbone were
assigned to the newly erected tyrannosaurid genus Zhuchengtyrannus in 2011. It is possible
that T. zhuchengensis is synonymous with Zhuchengtyrannus. In any case, T. zhuchengensis is
considered to be a nomen dubium as the holotype lacks diagnostic features below the level
Tyrannosaurinae.[78]

In 2006, a fragmentary tyrannosaurid lacrimal (CM 9401) from the Judith River Formation of
Fergus County, Montana was described as ?Tyrannosaurus sp. This isolated right lacrimal was
originally collected alongside the holotype specimen of Deinosuchus rugosus, a giant
crocodylian, and remained undescribed until its re-identification as belonging to a
tyrannosaurid theropod in the 1980s by paleontologist Dale Russell. The lacrimal closely
resembles those of Tyrannosaurus rex in both size and morphology. Notably, it lacks the
"lacrimal horn" typically present in earlier tyrannosaurids like Albertosaurus and Gorgosaurus,
instead exhibiting a distinct rugosity along the dorsal surface—consistent with T. rex and its
Asian relative Tarbosaurus. The specimen's considerable size places it within the range of
known T. rex individuals, suggesting the presence of large tyrannosaurids during the
Campanian stage (~75 million years ago), a temporal range earlier than the established
Maastrichtian age (~68–66 Ma) for Tyrannosaurus rex. However, the exact age and
provenance of CM 9401 remain uncertain due to a lack of detailed field documentation.[79]

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In a 2022 study, Gregory S. Paul and colleagues argued that Tyrannosaurus rex, as traditionally
understood, actually represents three species: the type species Tyrannosaurus rex, and two new
species: T. imperator (meaning "tyrant lizard emperor") and T. regina (meaning "tyrant lizard
queen"). The holotype of the former (T. imperator) is the Sue specimen, and the holotype of the
latter (T. regina) is Wankel rex. The division into multiple species was primarily based on the
observation of a very high degree of variation in the proportions and robusticity of the femur
(and other skeletal elements) across catalogued T. rex specimens, more so than that observed in
other theropods recognized as one species. Differences of general body proportions
representing robust and gracile morphotypes were also used as a line of evidence, in addition to
the number of small, slender incisiform teeth in the dentary, as based on tooth sockets.
Specifically, the paper's T. rex was distinguished by robust anatomy, a moderate ratio of femur
length vs circumference, and the possession of a singular slender incisiform dentary tooth; T.
imperator was considered to be robust with a small femur length to circumference ratio and
two of the slender teeth; and T. regina was a gracile form with a high femur ratio and one of the
slender teeth. It was observed that variation in proportions and robustness became more
extreme higher up in the sample, stratigraphically. This was interpreted as a single earlier
population, T. imperator, speciating into more than one taxon, T. rex and T. regina.[80]

However, several other leading paleontologists, including Stephen Brusatte, Thomas Carr,
Thomas Holtz, David Hone, Jingmai O'Connor, and Lindsay Zanno, criticized the study or
expressed skepticism of its conclusions when approached by various media outlets for
comment.[81][82][83] Their criticism was subsequently published in a technical paper.[84] Holtz
and Zanno both remarked that it was plausible that more than one species of Tyrannosaurus
existed, but felt the new study was insufficient to support the species it proposed. Holtz
remarked that, even if Tyrannosaurus imperator represented a distinct species from
Tyrannosaurus rex, it may represent the same species as Nanotyrannus lancensis and would
need to be called Tyrannosaurus lancensis. O'Connor, a curator at the Field Museum, where
the T. imperator holotype Sue is displayed, regarded the new species as too poorly-supported to
justify modifying the exhibit signs. Brusatte, Carr, and O'Connor viewed the distinguishing
features proposed between the species as reflecting natural variation within a species. Both Carr
and O'Connor expressed concerns about the study's inability to determine which of the
proposed species several well-preserved specimens belonged to. Another paleontologist, Philip
J. Currie, originally co-authored the study but withdrew from it as he did not want to be
involved in naming the new species.[81] Paul still rejected the objections raised by critics,
insisting that they are unwilling to consider that Tyrannosaurus might represent more than one
species.[85]

Tyrannosaurus mcraeensis
In 2024, Dalman and colleagues described the remains of a tyrannosaur discovered in 1983 in
the Campanian-early Maastrichtian Hall Lake Formation in New Mexico. Reposited at the New
Mexico Museum of Natural History and Science, the fossil material (NMMNH P-3698) consists
of the right postorbital, right squamosal, left palatine, and an incomplete maxilla from the skull,

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the left dentary, right splenial, right prearticular, right

angular and right articular from the lower jaws, isolated
teeth, and chevrons.[68] Some of the bones were briefly
mentioned in 1984 as belonging to T. rex,[86] and described
in 1986.[87]

Lehman and Carpenter (1990) suggested that NMMNH P-

3698 belonged to a new tyrannosaurid genus,[88] while Carr
and Williamson (2000) disagreed with their claim.[89]
Sullivan and Lucas (2015) argued that there is little evidence
to support NMMNH P-3698 as a specimen of
Tyrannosaurus rex, so they tentatively classified it as cf.
Tyrannosaurus sp.; they also considered that the McRae
tyrannosaur lived before the Lancian (before 67 million
years ago) based on its coexistence with Alamosaurus.[90] Reconstructed skull of T. mcraeensis

Dalman et al. (2024) proposed the new name

Tyrannosaurus mcraeensis for the holotype (NMMNH P-3698), referencing the McRae Group,
the rock layers to which the Hall Lake Formation belongs. These rock layers were estimated to
date to between 72.7 and 70.9 Ma, correlating to the latest Campanian or earliest
Maastrichtian.[68] U-Pb zircon age estimates by Schantz and Amato (2024) also support the late
Campanian to early Maastrichtian age of the Hall Lake Formation, with the mean estimate of
74.1 ± 0.9 Ma at 10 metres (33 ft) above the base of the formation and the maximum
depositional age of 69.8 ± 0.7 Ma based on a sandstone from this fossil locality.[91] The holotype
of T. mcraeensis is found in the strata that are around a few million years older than the
accepted range of T. rex, which existed at the end of the Maastrichtian. T. mcraeensis was
estimated at 12 metres (39 ft) long, which is similar to the size of an adult T. rex. The two are
distinguished by characters of the skull. Amongst these, the dentary of T. mcraeensis is
proportionately longer and possesses a less prominent chin, and the lower jaw shallower than
that of T. rex, suggesting a weaker bite. The teeth are likewise blunter and more laterally
compressed, while the post orbital crests are less prominent. Likewise, the skeletal anatomy
showcases shared characteristics with Tarbosaurus and Zhuchengtyrannus.[68][92]

Nanotyrannus
Other tyrannosaurid fossils found in the same formations as T. rex were originally classified as
separate taxa, including Aublysodon and Albertosaurus megagracilis,[62] the latter being
named Dinotyrannus megagracilis in 1995.[93] These fossils are now universally considered to
belong to juvenile T. rex.[94] A small but nearly complete skull from Montana, 60 centimeters
(2.0 ft) long, might be an exception. This skull, CMNH 7541, was originally classified as a
species of Gorgosaurus (G. lancensis) by Charles W. Gilmore in 1946.[95] In 1988, the specimen
was re-described by Robert T. Bakker, Phil Currie, and Michael Williams, then the curator of
paleontology at the Cleveland Museum of Natural History, where the original specimen was
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housed and is now on display. Their initial research

indicated that the skull bones were fused, and that it
therefore represented an adult specimen. In light of this,
Bakker and colleagues assigned the skull to a new genus
named Nanotyrannus (meaning "dwarf tyrant", for its
apparently small adult size). The specimen is estimated to
have been around 5.2 meters (17 ft) long when it died.[96]
However, In 1999, a detailed analysis by Thomas Carr Cast of CMNH 7541, the holotype of

revealed the specimen to be a juvenile, leading Carr and Nanotyrannus lancensis, sometimes
many other paleontologists to consider it a juvenile T. rex interpreted as a juvenile
individual.[97][98] Tyrannosaurus.

In 2001, a more
complete juvenile tyrannosaur (nicknamed "Jane", catalog
number BMRP 2002.4.1), belonging to the same species as
the original Nanotyrannus specimen, was uncovered. This
discovery prompted a conference on tyrannosaurs focused
on the issues of Nanotyrannus validity at the Burpee
Museum of Natural History in 2005. Several paleontologists
who had previously published opinions that N. lancensis
was a valid species, including Currie and Williams, saw the
Reconstructed skeleton of "Jane",
Burpee Museum of Natural History
discovery of "Jane" as a confirmation that Nanotyrannus
was, in fact, a juvenile T. rex.[99][100][101] Peter Larson
continued to support the hypothesis that N. lancensis was a
separate but closely related species, based on skull features such as two more teeth in both jaws
than T. rex; as well as proportionately larger hands with phalanges on the third metacarpal and
different wishbone anatomy in an undescribed specimen. He also argued that Stygivenator,
generally considered to be a juvenile T. rex, may be a younger Nanotyrannus
specimen.[102][103] Later research revealed that other tyrannosaurids such as Gorgosaurus also
experienced reduction in tooth count during growth,[97] and given the disparity in tooth count
between individuals of the same age group in this genus and Tyrannosaurus, this feature may
also be due to individual variation.[98] In 2013, Carr noted that all of the differences claimed to
support Nanotyrannus have turned out to be individually or ontogenetically variable features or
products of distortion of the bones.[104]

In 2016, analysis of limb proportions by Persons and Currie suggested Nanotyrannus

specimens to have differing cursoriality levels, potentially separating it from T. rex.[105]
However, paleontologist Manabu Sakomoto has commented that this conclusion may be
impacted by low sample size, and the discrepancy does not necessarily reflect taxonomic
distinction.[106] In 2016, Joshua Schmerge argued for Nanotyrannus' validity based on skull
features, including a dentary groove in BMRP 2002.4.1's skull. According to Schmerge, as that
feature is absent in T. rex and found only in Dryptosaurus and albertosaurines, this suggests
Nanotyrannus is a distinct taxon within the Albertosaurinae.[107] The same year, Carr and

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colleagues noted that this was insufficient to clarify

Nanotyrannus' validity or classification, being a common
and ontogenetically variable feature among
[108]
tyrannosauroids.

A 2020 study by Holly Woodward and colleagues showed

the specimens referred to Nanotyrannus were all
ontogenetically immature and found it probable that these
specimens belonged to T. rex.[109] The same year, Carr Adult T. rex skeleton (the specimen
AMNH 5027) at American Museum
published a paper on T. rex's growth history, finding that of Natural History.
CMNH 7541 fit within the expected ontogenetic variation of
the taxon and displayed juvenile characteristics found in
other specimens. It was classified as a juvenile, under 13 years old with a skull less than 80 cm
(31 in). No significant sexual or phylogenetic variation was discernible among any of the 44
specimens studied, with Carr stating that characters of potential phylogenetic importance
decrease throughout age at the same rate as growth occurs.[110] Discussing the paper's results,
Carr described how all Nanotyrannus specimens formed a continual growth transition between
the smallest juveniles and the subadults, unlike what would be expected if it were a distinct
taxon where the specimens would group to the exclusion of Tyrannosaurus. Carr concluded
that "the 'nanomorphs' are not all that similar to each other and instead form an important
bridge in the growth series of T. rex that captures the beginnings of the profound change from
the shallow skull of juveniles to the deep skull that is seen in fully-developed adults."[111]

However, a 2024 paper published by Nick Longrich and Evan Thomas Saitta reexamined the
holotype and referred specimens of Nanotyrannus. Based on several factors, including
differences in morphology, ontogeny, and phylogeny, Longrich and Saitta suggest that
Nanotyrannus is a distinct taxon which may fall outside of Tyrannosauridae, based on some of
their phylogenetic analyses.[112]

Paleobiology

Life history
The identification of several specimens as juvenile T. rex has allowed scientists to document
ontogenetic changes in the species, estimate the lifespan, and determine how quickly the
animals would have grown. The smallest known individual (LACM 28471, the "Jordan
theropod") is estimated to have weighed only 30 kg (66 lb), while the largest adults, such as
FMNH PR2081 (Sue) most likely weighed about 5,650 kg (12,460 lb). Histologic analysis of T.
rex bones showed LACM 28471 had aged only 2 years when it died, while Sue was 28 years old,
an age which may have been close to the maximum for the species.[40]

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Histology has also

allowed the age of other
specimens to be
determined. Growth
curves can be developed
when the ages of
different specimens are
plotted on a graph along Illustration of a juvenile
with their mass. A T. rex Tyrannosaurus rex
A graph showing the hypothesized growth curve is S-
growth curve, body mass versus age
shaped, with juveniles
(drawn in black, with other
tyrannosaurids for comparison).
remaining under 1,800 kg (4,000 lb) until approximately 14
Based on Erickson and colleagues years of age, when body size began to increase dramatically.
2004 During this rapid growth phase, a young T. rex would gain
an average of 600 kg (1,300 lb) a year for the next four
years. At 18 years of age, the curve plateaus again, indicating
that growth slowed dramatically. For example, only 600 kg (1,300 lb) separated the 28-year-old
Sue from a 22-year-old Canadian specimen (RTMP 81.12.1).[40] A 2004 histological study
performed by different workers corroborates these results, finding that rapid growth began to
slow at around 16 years of age.[113]

A study by Hutchinson and colleagues in 2011 corroborated the previous estimation methods in
general, but their estimation of peak growth rates is significantly higher; it found that the
"maximum growth rates for T. rex during the exponential stage are 1790 kg/year".[29] Although
these results were much higher than previous estimations, the authors noted that these results
significantly lowered the great difference between its actual growth rate and the one which
would be expected of an animal of its size.[29] The sudden change in growth rate at the end of
the growth spurt may indicate physical maturity, a hypothesis which is supported by the
discovery of medullary tissue in the femur of a 16 to 20-year-old T. rex from Montana (MOR
1125, also known as B-rex). Medullary tissue is found only in female birds during ovulation,
indicating that B-rex was of reproductive age.[114] Further study indicates an age of 18 for this
specimen.[115] In 2016, it was finally confirmed by Mary Higby Schweitzer and Lindsay Zanno
and colleagues that the soft tissue within the femur of MOR 1125 was medullary tissue. This also
confirmed the identity of the specimen as a female. The discovery of medullary bone tissue
within Tyrannosaurus may prove valuable in determining the sex of other dinosaur species in
future examinations, as the chemical makeup of medullary tissue is unmistakable.[116] Other
tyrannosaurids exhibit extremely similar growth curves, although with lower growth rates
corresponding to their lower adult sizes.[117]

An additional study published in 2020 by Woodward and colleagues, for the journal Science
Advances indicates that during their growth from juvenile to adult, Tyrannosaurus was capable
of slowing down its growth to counter environmental factors such as lack of food. The study,
focusing on two juvenile specimens between 13 and 15 years old housed at the Burpee Museum

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in Illinois, indicates that the rate of maturation for

Tyrannosaurus was dependent on resource abundance.
This study also indicates that in such changing
environments, Tyrannosaurus was particularly well-suited
to an environment that shifted yearly in regards to resource
abundance, hinting that other midsize predators might have
had difficulty surviving in such harsh conditions and
explaining the niche partitioning between juvenile and adult
tyrannosaurs. The study further indicates that
Tyrannosaurus and the dubious genus Nanotyrannus are
synonymous, due to analysis of the growth rings in the
bones of the two specimens studied.[118][119]

Over half of the known T. rex specimens appear to have died

within six years of reaching sexual maturity, a pattern which
is also seen in other tyrannosaurs and in some large, long-
lived birds and mammals today. These species are
characterized by high infant mortality rates, followed by Diagram showing growth stages
relatively low mortality among juveniles. Mortality increases
again following sexual maturity, partly due to the stresses of
reproduction. One study suggests that the rarity of juvenile T. rex fossils is due in part to low
juvenile mortality rates; the animals were not dying in large numbers at these ages, and thus
were not often fossilized. This rarity may also be due to the incompleteness of the fossil record
or to the bias of fossil collectors towards larger, more spectacular specimens.[117] In a 2013
lecture, Thomas Holtz Jr. suggested that dinosaurs "lived fast and died young" because they
reproduced quickly whereas mammals have long lifespans because they take longer to
reproduce.[120] Gregory S. Paul also writes that Tyrannosaurus reproduced quickly and died
young but attributes their short lifespans to the dangerous lives they lived.[121]

Skin and possible filamentous feathering

The discovery of feathered dinosaurs led to debate regarding
whether, and to what extent, Tyrannosaurus might have
been feathered.[122][123] Filamentous structures, which are
commonly recognized as the precursors of feathers, have
been reported in the small-bodied, basal tyrannosauroid
Dilong paradoxus from the Early Cretaceous Yixian
Formation of China in 2004.[124] Because integumentary
impressions of larger tyrannosauroids known at that time Fossilized skin impressions from the
showed evidence of scales, the researchers who studied tail region of a Tyrannosaurus rex,
Dilong speculated that insulating feathers might have been Houston Museum of Natural Science
lost by larger species due to their smaller surface-to-volume
ratio.[124] The subsequent discovery of the giant species

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Yutyrannus huali, also from the Yixian, showed that even some large tyrannosauroids had
feathers covering much of their bodies, casting doubt on the hypothesis that they were a size-
related feature.[125] A 2017 study reviewed known skin impressions of tyrannosaurids, including
those of a Tyrannosaurus specimen nicknamed "Wyrex" (HMNS 2006.1743.01, formerly
known as BHI 6230) which preserves patches of mosaic scales on the tail, hip, and neck.[122]
The study concluded that feather covering of large tyrannosaurids such as Tyrannosaurus was,
if present, limited to the upper side of the trunk.[122]

A conference abstract published in 2016 posited that theropods such as Tyrannosaurus had
their upper teeth covered in lips, instead of bare teeth as seen in crocodilians. This was based on
the presence of enamel, which according to the study needs to remain hydrated, an issue not
faced by aquatic animals like crocodilians.[57] However, there has been criticism where it favors
the idea for lips, with the 2017 analytical study proposing that tyrannosaurids had large, flat
scales on their snouts instead of lips, as modern crocodiles do.[55][126] But crocodiles possess
rather cracked keratinized skin, not flat scales; by observing the hummocky rugosity of
tyrannosaurids, and comparing it to extant lizards, researchers have found that tyrannosaurids
had squamose scales rather than a crocodillian-like skin.[127][128]

In 2023, Cullen and colleagues supported the idea that theropods like tyrannosaurids had lips
based on anatomical patterns, such as those of the foramina on their face and jaws, more
similar to those of modern squamates such as monitor lizards or marine iguanas than those of
modern crocodilians like alligators. Comparison of the teeth of Daspletosaurus and American
alligators shows that the enamel of tyrannosaurids had no significant wear and that the teeth of
modern crocodilians were eroded on the labial side and were substantially worn. This suggests
that it is likely that theropod teeth were kept wet by lips. On the basis of the relationship
between hydration and wear resistance, the authors argued that it is unlikely that the teeth of
theropods, including tyrannosaurids, would have remained unworn when exposed for a long
time, because it would have been hard to maintain hydration. The authors also performed
regression analyses to demonstrate the relationship between tooth height and skull length, and
found that varanids like the crocodile monitor had substantially greater ratios of tooth height to
skull length than those of Tyrannosaurus, indicating that the teeth of theropods were not too
big to be covered by extraoral tissues when the mouth was closed.[59]

Sexual dimorphism
As the number of known specimens increased, scientists began to analyze the variation between
individuals and discovered what appeared to be two distinct body types, or morphs, similar to
some other theropod species. As one of these morphs was more solidly built, it was termed the
'robust' morph while the other was termed 'gracile'. Several morphological differences
associated with the two morphs were used to analyze sexual dimorphism in T. rex, with the
'robust' morph usually suggested to be female. For example, the pelvis of several 'robust'
specimens seemed to be wider, perhaps to allow the passage of eggs.[129] It was also thought

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that the 'robust' morphology correlated with a reduced chevron on

the first tail vertebra, also ostensibly to allow eggs to pass out of the
reproductive tract, as had been erroneously reported for
crocodiles.[130]

In recent years, evidence for sexual dimorphism has been

weakened. A 2005 study reported that previous claims of sexual
dimorphism in crocodile chevron anatomy were in error, casting
doubt on the existence of similar dimorphism between T. rex
sexes.[131] A full-sized chevron was discovered on the first tail
vertebra of Sue, an extremely robust individual, indicating that this
feature could not be used to differentiate the two morphs anyway.
As T. rex specimens have been found from Saskatchewan to New
Mexico, differences between individuals may be indicative of
geographic variation rather than sexual dimorphism. The
differences could also be age-related, with 'robust' individuals being
older animals.[51]

Skeleton casts mounted in aOnly a single Tyrannosaurus specimen has been conclusively
mating position, Jurassic shown to belong to a specific sex. Examination of B-rex
Museum of Asturias
demonstrated the preservation of soft tissue within several bones.
Some of this tissue has been identified as a medullary tissue, a
specialized tissue grown only in modern birds as a source of calcium for the production of
eggshell during ovulation. As only female birds lay eggs, medullary tissue is only found naturally
in females, although males are capable of producing it when injected with female reproductive
hormones like estrogen. This strongly suggests that B-rex was female and that she died during
ovulation.[114] Recent research has shown that medullary tissue is never found in crocodiles,
which are thought to be the closest living relatives of dinosaurs. The shared presence of
medullary tissue in birds and other theropod dinosaurs is further evidence of the close
evolutionary relationship between the two.[132]

Posture
Like many bipedal dinosaurs, T. rex was historically depicted as a 'living tripod', with the body
at 45 degrees or less from the vertical and the tail dragging along the ground, similar to a
kangaroo. This concept dates from Joseph Leidy's 1865 reconstruction of Hadrosaurus, the first
to depict a dinosaur in a bipedal posture.[133] In 1915, convinced that the creature stood upright,
Henry Fairfield Osborn, former president of the American Museum of Natural History, further
reinforced the notion in unveiling the first complete T. rex skeleton arranged this way. It stood
in an upright pose for 77 years, until it was dismantled in 1992.[134]

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By 1970, scientists realized this pose was incorrect and could

not have been maintained by a living animal, as it would
have resulted in the dislocation or weakening of several
joints, including the hips and the articulation between the
head and the spinal column.[135] The inaccurate AMNH
mount inspired similar depictions in many films and
paintings (such as Rudolph Zallinger's famous mural The
Age of Reptiles in Yale University's Peabody Museum of
Natural History)[136] until the 1990s, when films such as Outdated reconstruction (by Charles
Jurassic Park introduced a more accurate posture to the R. Knight), showing upright pose

general public.[137] Modern representations in museums,

art, and film show T. rex with its body approximately parallel to the ground with the tail
extended behind the body to balance the head.[138]

To sit down, Tyrannosaurus may have settled its weight backwards and rested its weight on a
pubic boot, the wide expansion at the end of the pubis in some dinosaurs. With its weight rested
on the pelvis, it may have been free to move the hindlimbs. Getting back up again might have
involved some stabilization from the diminutive forelimbs.[139][135] The latter known as
Newman's pushup theory has been debated. Nonetheless, Tyrannosaurus was probably able to
get up if it fell, which only would have required placing the limbs below the center of gravity,
with the tail as an effective counterbalance. Healed stress fractures in the forelimbs have been
put forward both as evidence that the arms cannot have been very useful[140][141] and as
evidence that they were indeed used and acquired wounds,[142] like the rest of the body.

Arms
When T. rex was first discovered, the humerus was the only
element of the forelimb known.[6] For the initial mounted
skeleton as seen by the public in 1915, Osborn substituted
longer, three-fingered forelimbs like those of Allosaurus.[4]
A year earlier, Lawrence Lambe described the short, two-
fingered forelimbs of the closely related Gorgosaurus.[143]
This strongly suggested that T. rex had similar forelimbs,
The forelimbs might have been used
but this hypothesis was not confirmed until the first
to help T. rex rise from a resting complete T. rex forelimbs were identified in 1989, belonging
pose, as seen in this cast (Bucky to MOR 555 (the "Wankel rex").[144][145] The remains of Sue
specimen) also include complete forelimbs.[51] T. rex arms are very
small relative to overall body size, measuring only 1 meter
(3.3 ft) long, and some scholars have labelled them as vestigial. However, the bones show large
areas for muscle attachment, indicating considerable strength. This was recognized as early as
1906 by Osborn, who speculated that the forelimbs may have been used to grasp a mate during
copulation.[8] Newman (1970) suggested that the forelimbs were used to assist Tyrannosaurus

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in rising from a prone position.[135] Since then, other functions have been proposed, although
some scholars find them implausible.[141] Padian (2022) argued that the reduction of the arms
in tyrannosaurids did not serve a particular function but was a secondary adaptation, stating
that as tyrannosaurids developed larger and more powerful skulls and jaws, the arms got
smaller to avoid being bitten or torn by other individuals, particularly during group
feedings.[141]

Another possibility is that the forelimbs held struggling prey

while it was killed by the tyrannosaur's enormous jaws. This
hypothesis may be supported by biomechanical analysis. T.
rex forelimb bones exhibit extremely thick cortical bone,
which has been interpreted as evidence that they were
developed to withstand heavy loads. The biceps brachii
muscle of an adult T. rex was capable of lifting 199
kilograms (439 lb) by itself; other muscles such as the
brachialis would work along with the biceps to make elbow Diagram illustrating arm anatomy
flexion even more powerful. The M. biceps muscle of T. rex
was 3.5 times as powerful as the human equivalent. A T. rex
forearm had a limited range of motion, with the shoulder and elbow joints allowing only 40 and
45 degrees of motion, respectively. In contrast, the same two joints in Deinonychus allow up to
88 and 130 degrees of motion, respectively, while a human arm can rotate 360 degrees at the
shoulder and move through 165 degrees at the elbow. The heavy build of the arm bones,
strength of the muscles, and limited range of motion may indicate a system evolved to hold fast
despite the stresses of a struggling prey animal. In the first detailed scientific description of
Tyrannosaurus forelimbs, paleontologists Kenneth Carpenter and Matt Smith dismissed
notions that the forelimbs were useless or that Tyrannosaurus was an obligate scavenger.[146]

The idea that the arms served as weapons when hunting prey have also been proposed by
Steven M. Stanley, who suggested that the arms were used for slashing prey, especially by using
the claws to rapidly inflict long, deep gashes to its prey.[147] This was dismissed by Padian, who
argued that Stanley based his conclusion on incorrectly estimated forelimb size and range of
motion.[141]

Thermoregulation
Tyrannosaurus, like most dinosaurs, was long thought to
have an ectothermic ("cold-blooded") reptilian metabolism.
The idea of dinosaur ectothermy was challenged by
scientists like Robert T. Bakker and John Ostrom in the
Restoration showing partial early years of the "Dinosaur Renaissance", beginning in the
feathering late 1960s.[148][149] T. rex itself was claimed to have been
endothermic ("warm-blooded"), implying a very active
lifestyle.[38] Since then, several paleontologists have sought to determine the ability of

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Tyrannosaurus to regulate its body temperature. Histological evidence of high growth rates in
young T. rex, comparable to those of mammals and birds, may support the hypothesis of a high
metabolism. Growth curves indicate that, as in mammals and birds, T. rex growth was limited
mostly to immature animals, rather than the indeterminate growth seen in most other
vertebrates.[113]

Oxygen isotope ratios in fossilized bone are sometimes used to determine the temperature at
which the bone was deposited, as the ratio between certain isotopes correlates with
temperature. In one specimen, the isotope ratios in bones from different parts of the body
indicated a temperature difference of no more than 4 to 5 °C (7 to 9 °F) between the vertebrae of
the torso and the tibia of the lower leg. This small temperature range between the body core and
the extremities was claimed by paleontologist Reese Barrick and geochemist William Showers
to indicate that T. rex maintained a constant internal body temperature (homeothermy) and
that it enjoyed a metabolism somewhere between ectothermic reptiles and endothermic
mammals.[150] Other scientists have pointed out that the ratio of oxygen isotopes in the fossils
today does not necessarily represent the same ratio in the distant past, and may have been
altered during or after fossilization (diagenesis).[151] Barrick and Showers have defended their
conclusions in subsequent papers, finding similar results in another theropod dinosaur from a
different continent and tens of millions of years earlier in time (Giganotosaurus).[152]
Ornithischian dinosaurs also showed evidence of homeothermy, while varanid lizards from the
same formation did not.[153] In 2022, Wiemann and colleagues used a different approach—the
spectroscopy of lipoxidation signals, which are byproducts of oxidative phosphorylation and
correlate with metabolic rates—to show that various dinosaur genera including Tyrannosaurus
had endothermic metabolisms, on par with that of modern birds and higher than that of
mammals. They also suggested that such a metabolism was ancestrally common to all
dinosaurs.[154]

Even if T. rex does exhibit evidence of homeothermy, it does not necessarily mean that it was
endothermic. Such thermoregulation may also be explained by gigantothermy, as in some living
sea turtles.[155][156][157] Similar to contemporary crocodilians, openings (dorsotemporal
fenestrae) in the skull roofs of Tyrannosaurus may have aided thermoregulation.[158]

Soft tissue
In the March 2005 issue of Science, Mary Higby Schweitzer of North Carolina State University
and colleagues announced the recovery of soft tissue from the marrow cavity of a fossilized leg
bone from a T. rex. The bone had been intentionally, though reluctantly, broken for shipping
and then not preserved in the normal manner, specifically because Schweitzer was hoping to
test it for soft tissue.[159] Designated as the Museum of the Rockies specimen 1125, or MOR
1125, the dinosaur was previously excavated from the Hell Creek Formation. Flexible,
bifurcating blood vessels and fibrous but elastic bone matrix tissue were recognized. In
addition, microstructures resembling blood cells were found inside the matrix and vessels. The
structures bear resemblance to ostrich blood cells and vessels. Whether an unknown process,

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distinct from normal fossilization, preserved the material, or the

material is original, the researchers do not know, and they are
careful not to make any claims about preservation.[160] If it is found
to be original material, any surviving proteins may be used as a
means of indirectly guessing some of the DNA content of the
dinosaurs involved, because each protein is typically created by a
specific gene. The absence of previous finds may be the result of
people assuming preserved tissue was impossible, therefore not
looking. Since the first, two more tyrannosaurs and a hadrosaur
have also been found to have such tissue-like structures.[159] T. rex femur (MOR 1125)
Research on some of the tissues involved has suggested that birds from which demineralized
are closer relatives to tyrannosaurs than other modern animals.[161] matrix and peptides (insets)
were obtained
The original endogenous chemistry was also found in MOR 1125
based on preservation of elements associated with bone remodeling
and redeposition (sulfur, calcium, zinc), which showed that the bone cortices are similar to
those of extant birds.[162]

In studies reported in Science in April 2007, Asara and colleagues concluded that seven traces
of collagen proteins detected in purified T. rex bone most closely match those reported in
chickens, followed by frogs and newts. The discovery of proteins from a creature tens of millions
of years old, along with similar traces the team found in a mastodon bone at least 160,000 years
old, upends the conventional view of fossils and may shift paleontologists' focus from bone
hunting to biochemistry. Until these finds, most scientists presumed that fossilization replaced
all living tissue with inert minerals. Paleontologist Hans Larsson of McGill University in
Montreal, who was not part of the studies, called the finds "a milestone", and suggested that
dinosaurs could "enter the field of molecular biology and really slingshot paleontology into the
modern world".[163]

The presumed soft tissue was called into question by Thomas Kaye of the University of
Washington and his co-authors in 2008. They contend that what was really inside the
tyrannosaur bone was slimy biofilm created by bacteria that coated the voids once occupied by
blood vessels and cells.[164] The researchers found that what previously had been identified as
remnants of blood cells, because of the presence of iron, were actually framboids, microscopic
mineral spheres bearing iron. They found similar spheres in a variety of other fossils from
various periods, including an ammonite. In the ammonite, they found the spheres in a place
where the iron they contain could not have had any relationship to the presence of blood.[165]
Schweitzer has strongly criticized Kaye's claims and argues that there is no reported evidence
that biofilms can produce branching, hollow tubes like those noted in her study.[166] San
Antonio, Schweitzer and colleagues published an analysis in 2011 of what parts of the collagen
had been recovered, finding that it was the inner parts of the collagen coil that had been
preserved, as would have been expected from a long period of protein degradation.[167] Other
research challenges the identification of soft tissue as biofilm and confirms finding "branching,
vessel-like structures" from within fossilized bone.[168]

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Speed
Scientists have produced a wide range of possible maximum
Femur (thigh bone) running speeds for Tyrannosaurus: mostly around 9 meters
per second (32 km/h; 20 mph), but as low as 4.5–6.8 meters
per second (16–24 km/h; 10–15 mph) and as high as 20
Tibia (shin bone)
meters per second (72 km/h; 45 mph), though it running this
speed is very unlikely. Tyrannosaurus was a bulky and heavy
Metatarsals (foot bones) carnivore so it is unlikely to run very fast at all compared to
Dewclaw other theropods like Carnotaurus or Giganotosaurus.[169]
Phalanges (toe bones) Researchers have relied on various estimating techniques
Skeletal anatomy of a T. rex right because, while there are many tracks of large theropods
leg
walking, none showed evidence of running.[170]

A 2002 report used a mathematical model (validated by

applying it to three living animals: alligators, chickens, and humans; and eight more species,
including emus and ostriches[170]) to gauge the leg muscle mass needed for fast running (over
40 km/h or 25 mph).[169] Scientists who think that Tyrannosaurus was able to run point out
that hollow bones and other features that would have lightened its body may have kept adult
weight to a mere 4.5 metric tons (5.0 short tons) or so, or that other animals like ostriches and
horses with long, flexible legs are able to achieve high speeds through slower but longer
strides.[170] Proposed top speeds exceeded 40 kilometers per hour (25 mph) for
Tyrannosaurus, but were deemed infeasible because they would require exceptional leg
muscles of approximately 40–86% of total body mass. Even moderately fast speeds would have
required large leg muscles. If the muscle mass was less, only 18 kilometers per hour (11 mph) for
walking or jogging would have been possible.[169] Holtz noted that tyrannosaurids and some
closely related groups had significantly longer distal hindlimb components (shin plus foot plus
toes) relative to the femur length than most other theropods, and that tyrannosaurids and their
close relatives had a tightly interlocked metatarsus (foot bones).[171] The third metatarsal was
squeezed between the second and fourth metatarsals to form a single unit called an
arctometatarsus. This ankle feature may have helped the animal to run more efficiently.[172]
Together, these leg features allowed Tyrannosaurus to transmit locomotory forces from the foot
to the lower leg more effectively than in earlier theropods.[171]

Additionally, a 2020 study indicates that Tyrannosaurus and other tyrannosaurids were
exceptionally efficient walkers. Studies by Dececchi et al., compared the leg proportions, body
mass, and the gaits of more than 70 species of theropod dinosaurs including Tyrannosaurus
and its relatives. The research team then applied a variety of methods to estimate each
dinosaur's top speed when running as well as how much energy each dinosaur expended while
moving at more relaxed speeds such as when walking. Among smaller to medium-sized species
such as dromaeosaurids, longer legs appear to be an adaptation for faster running, in line with
previous results by other researchers. But for theropods weighing over 1,000 kg (2,200 lb), top
running speed is limited by body size, so longer legs instead were found to have correlated with

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low-energy walking. The results further indicate that smaller

theropods evolved long legs as a means to both aid in
hunting and escape from larger predators while larger
theropods that evolved long legs did so to reduce the energy
costs and increase foraging efficiency, as they were freed
from the demands of predation pressure due to their role as
apex predators. Compared to more basal groups of
theropods in the study, tyrannosaurs like Tyrannosaurus
itself showed a marked increase in foraging efficiency due to Only known tyrannosaurid trackway
reduced energy expenditures during hunting or scavenging.
(Bellatoripes fredlundi), from the
This in turn likely resulted in tyrannosaurs having a reduced
Wapiti Formation, British Columbia
need for hunting forays and requiring less food to sustain
themselves as a result. Additionally, the research, in
conjunction with studies that show tyrannosaurs were more agile than other large-bodied
theropods, indicates they were quite well-adapted to a long-distance stalking approach followed
by a quick burst of speed to go for the kill. Analogies can be noted between tyrannosaurids and
modern wolves as a result, supported by evidence that at least some tyrannosaurids were
hunting in group settings.[173][174]

A study published in 2021 by Pasha van Bijlert et al., calculated the preferred walking speed of
Tyrannosaurus, reporting a speed of 1.28 meters per second (4.6 km/h; 2.9 mph). While
walking, animals reduce their energy expenditure by choosing certain step rhythms at which
their body parts resonate. The same would have been true for dinosaurs, but previous studies
did not fully account for the impact the tail had on their walking speeds. According to the
authors, when a dinosaur walked, its tail would slightly sway up and down with each step as a
result of the interspinous ligaments suspending the tail. Like rubber bands, these ligaments
stored energy when they are stretched due to the swaying of the tail. Using a 3-D model of
Tyrannosaurus specimen Trix, muscles and ligaments were reconstructed to simulate the tail
movements. This results in a rhythmic, energy-efficient walking speed for Tyrannosaurus
similar to that seen in living animals such as humans, ostriches and giraffes.[175]

A 2017 study estimated the top running speed of Tyrannosaurus as 17 mph (27 km/h),
speculating that Tyrannosaurus exhausted its energy reserves long before reaching top speed,
resulting in a parabola-like relationship between size and speed.[176][177] Another 2017 study
hypothesized that an adult Tyrannosaurus was incapable of running due to high skeletal loads.
Using a calculated weight estimate of 7 tons, the model showed that speeds above 11 mph
(18 km/h) would have probably shattered the leg bones of Tyrannosaurus. The finding may
mean that running was also not possible for other giant theropod dinosaurs like
Giganotosaurus, Mapusaurus and Acrocanthosaurus.[178] However, studies by Eric Snively
and colleagues, published in 2019 indicate that Tyrannosaurus and other tyrannosaurids were
more maneuverable than allosauroids and other theropods of comparable size due to low
rotational inertia compared to their body mass combined with large leg muscles. As a result, it is
hypothesized that Tyrannosaurus was capable of making relatively quick turns and could likely

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pivot its body more quickly when close to its prey, or that while turning, the theropod could
"pirouette" on a single planted foot while the alternating leg was held out in a suspended swing
during a pursuit. The results of this study potentially could shed light on how agility could have
contributed to the success of tyrannosaurid evolution.[179]

Possible footprints
Rare fossil footprints and trackways found in New Mexico
and Wyoming that are assigned to the ichnogenus
Tyrannosauripus have been attributed to being made by
Tyrannosaurus, based on the stratigraphic age of the rocks
they are preserved in. The first specimen, found in 1994 was
described by Lockley and Hunt and consists of a single,
large footprint. Another pair of ichnofossils, described in Depiction of Tyrannosaurus rising
2021, show a large tyrannosaurid rising from a prone from the ground, based on fossil
position by rising up using its elbows in conjunction with tracks described in 2021.
the pads on their feet to stand. These two unique sets of
fossils were found in Ludlow, Colorado and Cimarron, New
Mexico.[180] Another ichnofossil described in 2018, perhaps belonging to a juvenile
Tyrannosaurus or the dubious genus Nanotyrannus was uncovered in the Lance Formation of
Wyoming. The trackway itself offers a rare glimpse into the walking speed of tyrannosaurids,
and the trackmaker is estimated to have been moving at a speed of 4.5–8.0 kilometers per hour
(2.8–5.0 mph), significantly faster than previously assumed for estimations of walking speed in
tyrannosaurids.[181][182]

Brain and senses

A study conducted by Lawrence Witmer and Ryan Ridgely of Ohio
University found that Tyrannosaurus shared the heightened
sensory abilities of other coelurosaurs, highlighting relatively rapid
and coordinated eye and head movements; an enhanced ability to
sense low frequency sounds, which would allow tyrannosaurs to
track prey movements from long distances; and an enhanced sense
of smell.[183] A study published by Kent Stevens concluded that
Tyrannosaurus had keen vision. By applying modified perimetry to
facial reconstructions of several dinosaurs including
Tyrannosaurus, the study found that Tyrannosaurus had a
binocular range of 55 degrees, surpassing that of modern hawks.
Stevens estimated that Tyrannosaurus had 13 times the visual
The eye-sockets faced
mainly forwards, giving it
acuity of a human and surpassed the visual acuity of an eagle,
good binocular vision (Sue which is 3.6 times that of a person. Stevens estimated a limiting far
specimen).

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point (that is, the distance at which an object can be seen as separate from the horizon) as far as
6 km (3.7 mi) away, which is greater than the 1.6 km (1 mi) that a human can see.[45][46][184]

Thomas Holtz Jr. would note that high depth perception of Tyrannosaurus may have been due
to the prey it had to hunt, noting that it had to hunt ceratopsians such as Triceratops,
ankylosaurs such as Ankylosaurus, and hadrosaurs. He would suggest that this made precision
more crucial for Tyrannosaurus enabling it to, "get in, get that blow in and take it down." In
contrast, Acrocanthosaurus had limited depth perception because they hunted large sauropods,
which were relatively rare during the time of Tyrannosaurus.[120]

Though no Tyrannosaurus sclerotic ring has been found, Kenneth Carpenter estimated its size
based on that of Gorgosaurus. The inferred sclerotic ring for the Stan specimen is ~7 cm
(2.8 in) in diameter with an internal aperture diameter of ~3.5 cm (1.4 in). Based on eye
proportions in living reptiles, this implies a pupil diameter of about 2.5 cm (0.98 in), an iris
diameter about that of the sclerotic ring, and an eyeball diameter of 11–12 cm (4.3–4.7 in).
Carpenter also estimated an eyeball depth of ~7.7–9.6 cm (3.0–3.8 in). Based on these
calculations, the f-number for Stan's eye is 3–3.8; since diurnal animals have f-numbers of 2.1
or higher, this would indicate that Tyrannosaurus had poor low-light vision and hunted during
the day.[185]

Tyrannosaurus had very large olfactory bulbs and olfactory nerves relative to their brain size,
the organs responsible for a heightened sense of smell. This suggests that the sense of smell was
highly developed, and implies that tyrannosaurs could detect carcasses by scent alone across
great distances. The sense of smell in tyrannosaurs may have been comparable to modern
vultures, which use scent to track carcasses for scavenging. Research on the olfactory bulbs has
shown that T. rex had the most highly developed sense of smell of 21 sampled non-avian
dinosaur species.[186]

Somewhat unusually among theropods, T. rex had a very

long cochlea. The length of the cochlea is often related to
hearing acuity, or at least the importance of hearing in
behavior, implying that hearing was a particularly important
sense to tyrannosaurs. Specifically, data suggests that T. rex
heard best in the low-frequency range, and that low-
frequency sounds were an important part of tyrannosaur Cast of the braincase at the
[183]
behavior. A 2017 study by Thomas Carr and colleagues Australian Museum, Sydney.
found that the snout of tyrannosaurids was highly sensitive,
based on a high number of small openings in the facial
bones of the related Daspletosaurus that contained sensory neurons. The study speculated that
tyrannosaurs might have used their sensitive snouts to measure the temperature of their nests
and to gently pick up eggs and hatchlings, as seen in modern crocodylians.[55] Another study
published in 2021 further suggests that Tyrannosaurus had an acute sense of touch, based on
neurovascular canals in the front of its jaws, which it could utilize to better detect and consume
prey. The study, published by Kawabe and Hittori et al., suggests that Tyrannosaurus could
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also accurately sense slight differences in material and movement, allowing it to utilize different
feeding strategies on different parts of its prey's carcasses depending on the situation. The
sensitive neurovascular canals of Tyrannosaurus also likely were adapted to performing fine
movements and behaviors such as nest building, parental care, and other social behavior such
as intraspecific communication. The results of this study also align with results made in
studying the related tyrannosaurid Daspletosaurus horneri and the allosauroid Neovenator,
which have similar neurovascular adaptations, suggesting that the faces of theropods were
highly sensitive to pressure and touch.[187][188] However, a more recent study reviewing the
evolution of the trigeminal canals among sauropsids notes that a much denser network of
neurovascular canals in the snout and lower jaw is more commonly encountered in aquatic or
semiaquatic taxa (e.g., Spinosaurus, Halszkaraptor, Plesiosaurus), and taxa that developed a
rhamphotheca (e.g., Caenagnathasia), while the network of canals in Tyrannosaurus appears
simpler, though still more derived than in most ornithischians, and overall terrestrial taxa such
as tyrannosaurids and Neovenator may have had average facial sensitivity for non-edentulous
terrestrial theropods, although further research is needed. The neurovascular canals in
Tyrannosaurus may instead have supported soft tissue structures for thermoregulation or
social signaling, the latter of which could be confirmed by the fact that the neurovascular
network of canals may have changed during ontogeny.[189]

A study by Grant R. Hurlburt, Ryan C. Ridgely and Lawrence Witmer obtained estimates for
Encephalization Quotients (EQs), based on reptiles and birds, as well as estimates for the ratio
of cerebrum to brain mass. The study concluded that Tyrannosaurus had the relatively largest
brain of all adult non-avian dinosaurs with the exception of certain small maniraptoriforms
(Bambiraptor, Troodon and Ornithomimus). The study found that Tyrannosaurus's relative
brain size was still within the range of modern reptiles, being at most 2 standard deviations
above the mean of non-avian reptile EQs. The estimates for the ratio of cerebrum mass to brain
mass would range from 47.5 to 49.53 percent. According to the study, this is more than the
lowest estimates for extant birds (44.6 percent), but still close to the typical ratios of the
smallest sexually mature alligators which range from 45.9–47.9 percent.[190] Other studies,
such as those by Steve Brusatte, indicate the encephalization quotient of Tyrannosaurus was
similar in range (2.0–2.4) to a chimpanzee (2.2–2.5), though this may be debatable as reptilian
and mammalian encephalization quotients are not equivalent.[191]

Social behavior
Philip J. Currie suggested that Tyrannosaurus may have been pack hunters, comparing T. rex to
related species Tarbosaurus bataar and Albertosaurus sarcophagus, citing fossil evidence that
may indicate gregarious (describing animals that travel in herds or packs) behavior.[192] A find
in South Dakota where three T. rex skeletons were in close proximity may suggest the formation
of a pack.[193][194] Cooperative pack hunting may have been an effective strategy for subduing
prey with advanced anti-predator adaptations which pose potential lethality such as Triceratops
and Ankylosaurus.[192]

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Currie's pack-hunting T. rex hypothesis has been criticized

for not having been peer-reviewed, but rather was discussed
in a television interview and book called Dino Gangs.[195]
The Currie theory for pack hunting by T. rex is based mainly
by analogy to a different species, Tarbosaurus bataar.
Evidence of gregariousness in T. bataar itself has not been
peer-reviewed, and to Currie's own admission, can only be
Mounted skeletons of different age
interpreted with reference to evidence in other closely
related species. According to Currie gregariousness in
groups (skeleton in lower left based
Albertosaurus sarcophagus is supported by the discovery of
on the juvenile formerly named
Stygivenator), Natural History
26 individuals with varied ages in the Dry Island bonebed.
Museum of Los Angeles County He ruled out the possibility of a predator trap due to the
similar preservation state of individuals and the near
absence of herbivores.[195][196]

Additional support of tyrannosaurid gregariousness can be found in fossilized trackways from

the Upper Cretaceous Wapiti Formation of northeastern British Columbia, Canada, left by three
tyrannosaurids traveling in the same direction.[197][198] According to scientists assessing the
Dino Gangs program, the evidence for pack hunting in Tarbosaurus and Albertosaurus is weak
and based on group skeletal remains for which alternate explanations may apply (such as
drought or a flood forcing dinosaurs to die together in one place).[195] Others researchers have
speculated that instead of large theropod social groups, some of these finds represent behavior
more akin to Komodo dragon-like mobbing of carcasses, even going as far as to say true pack-
hunting behavior may not exist in any non-avian dinosaurs due to its rarity in modern
predators.[199]

Evidence of intraspecific attack was found by Joseph Peterson and his colleagues in the juvenile
Tyrannosaurus nicknamed Jane. Peterson and his team found that Jane's skull showed healed
puncture wounds on the upper jaw and snout which they believe came from another juvenile
Tyrannosaurus. Subsequent CT scans of Jane's skull would further confirm the team's
hypothesis, showing that the puncture wounds came from a traumatic injury and that there was
subsequent healing.[200] The team would also state that Jane's injuries were structurally
different from the parasite-induced lesions found in Sue and that Jane's injuries were on its face
whereas the parasite that infected Sue caused lesions to the lower jaw.[201] Pathologies of other
Tyrannosaurus specimens have been suggested as evidence of conspecific attack, including
"Wyrex" with a hole penetrating its jugual and severe trauma on its tail that shows signs of bone
remodeling (not regrowth).[202][203]

Feeding strategies
Most paleontologists accept that Tyrannosaurus was both an active predator and a scavenger
like most large carnivores.[204] By far the largest carnivore in its environment, T. rex was most
likely an apex predator, preying upon hadrosaurs, armored herbivores like ceratopsians and

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ankylosaurs, and possibly sauropods.[205] Enamel δ44/42Ca values

also suggest the possibility that T. rex occasionally fed on carcasses
of marine reptiles and fish washed up on the shores of the Western
Interior Seaway.[206] A study in 2012 by Karl Bates and Peter
Falkingham found that Tyrannosaurus had the most powerful bite
of any terrestrial animal that has ever lived, finding an adult
Tyrannosaurus could have exerted 35,000 to 57,000 N (7,868 to
12,814 lbf) of force in the back teeth.[207][208][209] Even higher
estimates were made by Mason B. Meers in 2003.[48] This allowed
it to crush bones during repetitive biting and fully consume the
carcasses of large dinosaurs.[21] Stephan Lautenschlager and
colleagues calculated that Tyrannosaurus was capable of a
maximum jaw gape of around 80 degrees, a necessary adaptation
for a wide range of jaw angles to power the creature's strong
bite.[210][211]

A debate exists, however, about whether Tyrannosaurus was

primarily a predator or a pure scavenger. The debate originated in a
Tyrannosaurus tooth marks
1917 study by Lambe which argued that large theropods were pure
on bones of various
scavengers because Gorgosaurus teeth showed hardly any herbivorous dinosaurs
wear.[212] This argument disregarded the fact that theropods
replaced their teeth quite rapidly. Ever since the first
discovery of Tyrannosaurus most scientists have speculated
that it was a predator; like modern large predators it would
readily scavenge or steal another predator's kill if it had the
opportunity.[213]

Paleontologist Jack Horner has been a major proponent of

the view that Tyrannosaurus was not a predator at all but
instead was exclusively a scavenger.[144][214][215] He has put
forward arguments in the popular literature to support the A Tyrannosaurus mounted next to a
pure scavenger hypothesis: Triceratops at the Los Angeles
Natural History Museum
Tyrannosaur arms are short when compared to other
known predators. Horner argues that the arms were too
short to make the necessary gripping force to hold on to prey.[216] Other paleontologists
such as Thomas Holtz Jr. argued that there are plenty of modern-day predators that do not
use their forelimbs to hunt such as wolves, hyenas, and secretary birds as well as other
extinct animals thought to be predators that would not have used their forelimbs such as
phorusrhacids.[217][218]
Tyrannosaurs had large olfactory bulbs and olfactory nerves (relative to their brain size).
These suggest a highly developed sense of smell which could sniff out carcasses over great
distances, as modern vultures do. Research on the olfactory bulbs of dinosaurs has shown
that Tyrannosaurus had the most highly developed sense of smell of 21 sampled

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dinosaurs.[186]
Tyrannosaur teeth could crush bone, and therefore could extract as much food (bone
marrow) as possible from carcass remnants, usually the least nutritious parts. Karen Chin
and colleagues have found bone fragments in coprolites (fossilized feces) that they attribute
to tyrannosaurs, but point out that a tyrannosaur's teeth were not well adapted to
systematically chewing bone like hyenas do to extract marrow.[219]
Since at least some of Tyrannosaurus's potential prey could move quickly, evidence that it
walked instead of ran could indicate that it was a scavenger.[214] On the other hand, recent
analyses suggest that Tyrannosaurus, while slower than large modern terrestrial predators,
may well have been fast enough to prey on large hadrosaurs and ceratopsians.[169][24]
Other evidence suggests hunting behavior in Tyrannosaurus. The eye sockets of tyrannosaurs
are positioned so that the eyes would point forward, giving them binocular vision slightly better
than that of modern hawks. It is not obvious why natural selection would have favored this
long-term trend if tyrannosaurs had been pure scavengers, which would not have needed the
advanced depth perception that stereoscopic vision provides.[45][46] In modern animals,
binocular vision is found mainly in predators.

A skeleton of the hadrosaurid Edmontosaurus annectens

has been described from Montana with healed tyrannosaur-
inflicted damage on its tail vertebrae. The fact that the
damage seems to have healed suggests that the
Edmontosaurus survived a tyrannosaur's attack on a living
target, i.e. the tyrannosaur had attempted active
predation.[220] Despite the consensus that the tail bites were
caused by Tyrannosaurus, there has been some evidence to
The damage to the tail vertebrae of
show that they might have been created by other factors. For
this Edmontosaurus annectens
skeleton (on display at the Denver
example, a 2014 study suggested that the tail injuries might
Museum of Nature and Science) have been due to Edmontosaurus individuals stepping on
indicates that it may have been bitten each other,[221] while another study in 2020 backs up the
by a Tyrannosaurus hypothesis that biomechanical stress is the cause for the tail
injuries.[222] There is also evidence for an aggressive
interaction between a Triceratops and a Tyrannosaurus in the form of partially healed
tyrannosaur tooth marks on a Triceratops brow horn and squamosal (a bone of the neck frill);
the bitten horn is also broken, with new bone growth after the break. It is not known what the
exact nature of the interaction was, though: either animal could have been the aggressor.[223]
Since the Triceratops wounds healed, it is most likely that the Triceratops survived the
encounter and managed to overcome the Tyrannosaurus. In a battle against a bull Triceratops,
the Triceratops would likely defend itself by inflicting fatal wounds to the Tyrannosaurus using
its sharp horns.[224] Studies of Sue found a broken and healed fibula and tail vertebrae, scarred
facial bones and a tooth from another Tyrannosaurus embedded in a neck vertebra, providing
evidence for aggressive behavior.[225] Studies on hadrosaur vertebrae from the Hell Creek
Formation that were punctured by the teeth of what appears to be a late-stage juvenile

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Tyrannosaurus indicate that despite lacking the bone-crushing adaptations of the adults, young
individuals were still capable of using the same bone-puncturing feeding technique as their
adult counterparts.[226]

Tyrannosaurus may have had infectious saliva used to kill its prey, as proposed by William
Abler in 1992. Abler observed that the serrations (tiny protuberances) on the cutting edges of
the teeth are closely spaced, enclosing little chambers. These chambers might have trapped
pieces of carcass with bacteria, giving Tyrannosaurus a deadly, infectious bite much like the
Komodo dragon was thought to have.[227][228] Jack Horner and Don Lessem, in a 1993 popular
book, questioned Abler's hypothesis, arguing that Tyrannosaurus's tooth serrations as more
like cubes in shape than the serrations on a Komodo monitor's teeth, which are
rounded.[144]: 214–215

Tyrannosaurus, and most other theropods, probably primarily processed carcasses with lateral
shakes of the head, like crocodilians. The head was not as maneuverable as the skulls of
allosauroids, due to flat joints of the neck vertebrae.[229]

Cannibalism
Evidence also strongly suggests that tyrannosaurs were at least occasionally cannibalistic.
Tyrannosaurus itself has strong evidence pointing towards it having been cannibalistic in at
least a scavenging capacity based on tooth marks on the foot bones, humerus, and metatarsals
of one specimen.[230] Fossils from the Fruitland Formation, Kirtland Formation (both
Campanian in age) and the Maastrichtian aged Ojo Alamo Formation suggest that cannibalism
was present in various tyrannosaurid genera of the San Juan Basin. The evidence gathered from
the specimens suggests opportunistic feeding behavior in tyrannosaurids that cannibalized
members of their own species.[231] A study from Currie, Horner, Erickson and Longrich in 2010
has been put forward as evidence of cannibalism in the genus Tyrannosaurus.[230] They
studied some Tyrannosaurus specimens with tooth marks in the bones, attributable to the
same genus. The tooth marks were identified in the humerus, foot bones and metatarsals, and
this was seen as evidence for opportunistic scavenging, rather than wounds caused by
intraspecific combat. In a fight, they proposed it would be difficult to reach down to bite in the
feet of a rival, making it more likely that the bitemarks were made in a carcass. As the bitemarks
were made in body parts with relatively scantly amounts of flesh, it is suggested that the
Tyrannosaurus was feeding on a cadaver in which the more fleshy parts already had been
consumed. They were also open to the possibility that other tyrannosaurids practiced
cannibalism.[230]

Parenting
While there is no direct evidence of Tyrannosaurus raising their young (the rarity of juvenile
and nest Tyrannosaur fossils has left researchers guessing), it has been suggested by some that
like its closest living relatives, modern archosaurs (birds and crocodiles) Tyrannosaurus may

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have protected and fed its young. Crocodilians and birds are often suggested by some
paleontologists to be modern analogues for dinosaur parenting.[232] Direct evidence of parental
behavior exists in other dinosaurs such as Maiasaura peeblesorum, the first dinosaur to have
been discovered to raise its young, as well as more closely related Oviraptorids, the latter
suggesting parental behavior in theropods.[233][234][235][236][237]

Pathology
In 2001, Bruce Rothschild and others published a study
examining evidence for stress fractures and tendon
avulsions in 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.
Of the 81 Tyrannosaurus foot bones examined in the study,
one was found to have a stress fracture, while none of the 10
hand bones were found to have stress fractures. The Restoration of an individual (based
researchers found tendon avulsions only among on MOR 980) with parasite infections
Tyrannosaurus and Allosaurus. An avulsion injury left a
divot on the humerus of Sue the T. rex, apparently located at
the origin of the deltoid or teres major muscles. The presence of avulsion injuries being limited
to the forelimb and shoulder in both Tyrannosaurus and Allosaurus suggests that theropods
may have had a musculature more complex than and functionally different from those of birds.
The researchers concluded that Sue's tendon avulsion was probably obtained from struggling
prey. The presence of stress fractures and tendon avulsions, in general, provides evidence for a
"very active" predation-based diet rather than obligate scavenging.[238]

A 2009 study showed that smooth-edged holes in the skulls of several specimens might have
been caused by Trichomonas-like parasites that commonly infect birds. According to the study,
seriously infected individuals, including "Sue" and MOR 980 ("Peck's Rex"), might therefore
have died from starvation after feeding became increasingly difficult. Previously, these holes
had been explained by the bacterious bone infection Actinomycosis or by intraspecific
attacks.[239] A subsequent study found that while trichomoniasis has many attributes of the
model proposed (osteolytic, intra oral) several features make the assumption that it was the
cause of death less supportable by evidence. For example, the observed sharp margins with little
reactive bone shown by the radiographs of Trichomonas-infected birds are dissimilar to the
reactive bone seen in the affected T. rex specimens. Also, trichomoniasis can be very rapidly
fatal in birds (14 days or less) albeit in its milder form, and this suggests that if a Trichomonas-
like protozoan is the culprit, trichomoniasis was less acute in its non-avian dinosaur form
during the Late Cretaceous. Finally, the relative size of this type of lesions is much larger in
small bird throats, and may not have been enough to choke a T. rex.[240] A more recent study
examining the pathologies concluded that the osseous alteration observed most closely

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resembles those around healing human cranial trepanations and healing fractures in the
Triassic reptile Stagonolepis, in the absence of infection. The possible cause may instead have
been intraspecific combat.[241]

One study of Tyrannosaurus specimens with tooth marks in the bones attributable to the same
genus was presented as evidence of cannibalism.[230] Tooth marks in the humerus, foot bones
and metatarsals, may indicate opportunistic scavenging, rather than wounds caused by combat
with another T. rex.[230][242] Other tyrannosaurids may also have practiced cannibalism.[230]

Paleoecology
Tyrannosaurus lived during what is referred to
as the Lancian faunal stage (Maastrichtian age)
at the end of the Late Cretaceous.
Tyrannosaurus ranged from Canada in the
Fauna of Hell Creek (Tyrannosaurus in dark red, north to at least New Mexico in the south of
left). Laramidia.[5] During this time Triceratops was
the major herbivore in the northern portion of
its range, while the titanosaurian sauropod
Alamosaurus "dominated" its southern range. Tyrannosaurus remains have been discovered in
different ecosystems, including inland and coastal subtropical, and semi-arid plains.

Several notable Tyrannosaurus remains have been found in

the Hell Creek Formation. During the Maastrichtian this
area was subtropical, with a warm and humid climate. The
flora consisted mostly of angiosperms, but also included
trees like dawn redwood (Metasequoia) and Araucaria.
Tyrannosaurus shared this ecosystem with ceratopsians
Leptoceratops, Torosaurus, and Triceratops, the
hadrosaurid Edmontosaurus annectens, the parksosaurid Tyrannosaurus and other animals of
the Hell Creek Formation
Thescelosaurus, the ankylosaurs Ankylosaurus and
Denversaurus, the pachycephalosaurs Pachycephalosaurus
and Sphaerotholus, and the theropods Ornithomimus, Struthiomimus, Acheroraptor,
Dakotaraptor, Pectinodon and Anzu.[243]

Another formation with Tyrannosaurus remains is the Lance Formation of Wyoming. This has
been interpreted as a bayou environment similar to today's Gulf Coast. The fauna was very
similar to Hell Creek, but with Struthiomimus replacing its relative Ornithomimus. The small
ceratopsian Leptoceratops also lived in the area.[244]

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In its southern range, specifically based on remains discovered from the North Horn Formation
of Utah, Tyrannosaurus rex lived alongside the titanosaur Alamosaurus, the ceratopsid
Torosaurus and the indeterminate troodontids and hadrosaurids.[245][246] Tyrannosaurus
mcraeensis from the McRae Group of New Mexico coexisted with the ceratopsid Sierraceratops
and possibly the titanosaur Alamosaurus.[68] Potential remains identified as cf. Tyrannosaurus
have also been discovered from the Javelina Formation of Texas,[68] where the remains of the
titanosaur Alamosaurus, the ceratopsid Bravoceratops, the pterosaurs Quetzalcoatlus and
Wellnhopterus, and possible species of troodontids and hadrosaurids are found.[247][248][249]
Its southern range is thought to have been dominated by semi-arid inland plains, following the
probable retreat of the Western Interior Seaway as global sea levels fell.[250]

Tyrannosaurus may have also inhabited Mexico's Lomas Coloradas Formation in Sonora.
Though skeletal evidence is lacking, six shed and broken teeth from the fossil bed have been
thoroughly compared with other theropod genera and appear to be identical to those of
Tyrannosaurus. If true, the evidence indicates the range of Tyrannosaurus was possibly more
extensive than previously believed.[251] It is possible that tyrannosaurs were originally Asian
species, migrating to North America before the end of the Cretaceous period.[252]

Population estimates
According to studies published in 2021 by Charles Marshall
et al., the total population of adult Tyrannosaurus at any
given time was perhaps 20,000 individuals, with computer
estimations also suggesting a total population no lower than
1,300 and no higher than 328,000. The authors themselves
suggest that the estimate of 20,000 individuals is probably Chart of the time-averaged census
lower than what should be expected, especially when for large-bodied dinosaurs from the
factoring in that disease pandemics could easily wipe out entire Hell Creek Formation in the
study area
such a small population. Over the span of the genus'
existence, it is estimated that there were about 127,000
generations and that this added up to a total of roughly 2.5 billion animals until their
extinction.[253][254]

In the same paper, it is suggested that in a population of Tyrannosaurus adults numbering

20,000, the number of individuals living in an area the size of California could be as high as
3,800 animals, while an area the size of Washington D.C. could support a population of only
two adult Tyrannosaurus. The study does not take into account the number of juvenile animals
in the genus present in this population estimate due to their occupation of a different niche than
the adults, and thus it is likely the total population was much higher when accounting for this
factor. Simultaneously, studies of living carnivores suggest that some predator populations are
higher in density than others of similar weight (such as jaguars and hyenas, which are similar in
weight but have vastly differing population densities). Lastly, the study suggests that in most

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cases, only one in 80 million Tyrannosaurus would become fossilized, while the chances were
likely as high as one in every 16,000 of an individual becoming fossilized in areas that had more
dense populations.[253][254]

Meiri (2022) questioned the reliability of the estimates, citing uncertainty in metabolic rate,
body size, sex and age-specific survival rates, habitat requirements and range size variability as
shortcomings Marshall et al. did not take into account.[255] The authors of the original
publication replied that while they agree that their reported uncertainties were probably too
small, their framework is flexible enough to accommodate uncerainty in physiology, and that
their calculations do not depend on short-term changes in population density and geographic
range, but rather on their long-term averages. Finally, they remark that they did estimate the
range of reasonable survivorship curves and that they did include uncertainty in the time of
onset of sexual maturity and in the growth curve by incorporating the uncertainty in the
maximum body mass.[256]

Cultural significance
Since it was first described in 1905, T. rex has become the most widely recognized dinosaur
species in popular culture. It is the only dinosaur that is commonly known to the general public
by its full scientific name (binomial name) and the scientific abbreviation T. rex has also come
into wide usage.[51] Robert T. Bakker notes this in The Dinosaur Heresies and explains that, "a
name like 'T. rex' is just irresistible to the tongue."[38]

See also
History of paleontology
Sue (dinosaur) (FMNH-PR-2081)
Tyrannosauridae

Notes
a. lit. 'tyrant lizard'; from Ancient Greek τύραννος (túrannos) 'tyrant' and σαῦρος
(saûros) 'lizard'

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Further reading
Farlow, J. O.; Gatesy, S. M.; Holtz, T. R. Jr.; Hutchinson, J. R.; Robinson, J. M. (2000).
"Theropod Locomotion" (https://doi.org/10.1093%2Ficb%2F40.4.640). American Zoologist.
40 (4): 640–663. doi:10.1093/icb/40.4.640 (https://doi.org/10.1093%2Ficb%2F40.4.640).
JSTOR 3884284 (https://www.jstor.org/stable/3884284).

External links
The University of Edinburgh Lecture Dr Stephen Brusatte – Tyrannosaur Discoveries Feb
20, 2015 (https://www.youtube.com/watch?v=hVJmPmb_LWY)
28 species in the tyrannosaur family tree, when and where they lived (http://www.livescience
.com/53877-t-rex-was-invasive-species.html) Stephen Brusatte Thomas Carr 2016
Australia's answer to T-Rex (https://onesearch.slq.qld.gov.au/permalink/61SLQ_INST/dls06
p/alma99274923402061), State Library of Queensland

Exhibits
American Museum of Natural History (http://www.amnh.org/exhibitions/permanent-exhibition
s/fossil-halls/hall-of-saurischian-dinosaurs/tyrannosaurus-rex)

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