Tyrannosaurus Rex
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''Tyrannosaurus'' () is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of large theropod
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
. The
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
''Tyrannosaurus rex'' ( meaning 'king' in
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
), 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 North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
, on what was then an island continent known as
Laramidia Laramidia was an island continent that existed during the Late Cretaceous period (99.6–66 Year#SI prefix multipliers, Ma), when the Western Interior Seaway split the continent of North America in two. In the Mesozoic era, Laramidia was an island ...
. ''Tyrannosaurus'' had a much wider range than other tyrannosaurids.
Fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...
s are found in a variety of geological formations dating to the latest
Campanian The Campanian is the fifth of six ages of the Late Cretaceous epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campa ...
-
Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
ages of the late
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
period, 72.7 to 66 
million years ago Million years ago, abbreviated as Mya, Myr (megayear) or Ma (megaannum), is a unit of time equal to (i.e. years), or approximately 31.6 teraseconds. Usage Myr is in common use in fields such as Earth science and cosmology. Myr is also used w ...
, with isolated specimens possibly indicating an earlier origin in the middle Campanian. It was the last known member of the tyrannosaurids and among the last non- avian dinosaurs to exist before the
Cretaceous–Paleogene extinction event The Cretaceous–Paleogene (K–Pg) extinction event, also known as the K–T extinction, was the extinction event, mass extinction of three-quarters of the plant and animal species on Earth approximately 66 million years ago. The event cau ...
. Like other tyrannosaurids, ''Tyrannosaurus'' was a bipedal
carnivore A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they ar ...
with a massive skull balanced by a long, heavy tail. Relative to its large and powerful hind limbs, the forelimbs of ''Tyrannosaurus'' were short but unusually powerful for their size, and they had two clawed digits. The most complete specimen measures in length, but according to most modern estimates, ''Tyrannosaurus'' could have exceeded sizes of in length, in hip height, and in mass. Although some other theropods might have rivaled or exceeded ''Tyrannosaurus'' in
size Size in general is the Magnitude (mathematics), magnitude or dimensions of a thing. More specifically, ''geometrical size'' (or ''spatial size'') can refer to three geometrical measures: length, area, or volume. Length can be generalized ...
, it is still among the largest known land predators, with its estimated bite force being the largest among all terrestrial animals. By far the largest carnivore in its environment, ''Tyrannosaurus rex'' was most likely an
apex predator An apex predator, also known as a top predator or superpredator, is a predator at the top of a food chain, without natural predators of its own. Apex predators are usually defined in terms of trophic dynamics, meaning that they occupy the hig ...
, preying upon hadrosaurs, juvenile armored herbivores like
ceratopsia Ceratopsia or Ceratopia ( or ; Ancient Greek, Greek: "horned faces") is a group of herbivore, herbivorous, beaked dinosaurs that thrived in what are now North America, Asia and Europe, during the Cretaceous Period (geology), Period, although ance ...
ns and ankylosaurs, and possibly
sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their b ...
s. Some experts have suggested the dinosaur was primarily a
scavenger Scavengers are animals that consume Corpse decomposition, dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a he ...
. The question of whether ''Tyrannosaurus'' was an apex predator or a pure scavenger was among the longest debates in
paleontology Paleontology, also spelled as palaeontology or palæontology, is the scientific study of the life of the past, mainly but not exclusively through the study of fossils. Paleontologists use fossils as a means to classify organisms, measure ge ...
. Most paleontologists today accept that ''Tyrannosaurus'' was both a predator and a scavenger. Some specimens of ''Tyrannosaurus rex'' are nearly complete skeletons.
Soft tissue Soft tissue connective tissue, connects and surrounds or supports internal organs and bones, and includes muscle, tendons, ligaments, Adipose tissue, fat, fibrous tissue, Lymphatic vessel, lymph and blood vessels, fasciae, and synovial membranes. ...
and
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residue (biochemistry), residues. Proteins perform a vast array of functions within organisms, including Enzyme catalysis, catalysing metab ...
s have been reported in at least one of these specimens. The abundance of fossil material has allowed significant research into many aspects of the animal's biology, including its life history and
biomechanics Biomechanics is the study of the structure, function and motion of the mechanical aspects of biological systems, at any level from whole organisms to Organ (anatomy), organs, Cell (biology), cells and cell organelles, using the methods of mechani ...
. The feeding habits,
physiology Physiology (; ) is the science, scientific study of function (biology), functions and mechanism (biology), mechanisms in a life, living system. As a branches of science, subdiscipline of biology, physiology focuses on how organisms, organ syst ...
, and potential speed of ''Tyrannosaurus rex'' are a few subjects of debate. Its
taxonomy image:Hierarchical clustering diagram.png, 280px, Generalized scheme of taxonomy Taxonomy is a practice and science concerned with classification or categorization. Typically, there are two parts to it: the development of an underlying scheme o ...
is also controversial. The Asian '' Tarbosaurus bataar'' is very closely related to ''Tyrannosaurus'' and has sometimes been seen as a species of this genus. Several North American tyrannosaurids have been synonymized with ''Tyrannosaurus'', while some ''Tyrannosaurus'' specimens have been proposed as distinct species. The validity of these species, such as the more recently discovered ''T. mcraeensis'', is contentious. As the archetypal theropod, ''Tyrannosaurus'' has been one of the best-known dinosaurs since the early 20th century and has been featured in film, advertising, postal stamps, and many other media.


History of research


Earliest finds

A tooth from what is now documented as a ''Tyrannosaurus rex'' was found in July 1874 upon South Table Mountain (Colorado) by Jarvis Hall (Colorado) student Peter T. Dotson under the auspices of Prof. Arthur Lakes near Golden, Colorado. In the early 1890s, John Bell Hatcher collected postcranial elements in eastern
Wyoming Wyoming ( ) is a landlocked U.S. state, state in the Mountain states, Mountain West subregion of the Western United States, Western United States. It borders Montana to the north and northwest, South Dakota and Nebraska to the east, Idaho t ...
. The fossils were believed to be from the large species '' Ornithomimus grandis'' (now '' Deinodon'') but are now considered ''T. rex'' remains. In 1892,
Edward Drinker Cope Edward Drinker Cope (July 28, 1840 – April 12, 1897) was an American zoologist, paleontology, paleontologist, comparative anatomy, comparative anatomist, herpetology, herpetologist, and ichthyology, ichthyologist. Born to a wealthy Quaker fam ...
found two vertebral fragments of a large dinosaur. Cope believed the fragments belonged to an "agathaumid" ( ceratopsid) dinosaur, and named them ''Manospondylus gigas'', meaning "giant porous vertebra", in reference to the numerous openings for blood vessels he found in the bone. The ''M. gigas'' remains were, in 1907, identified by Hatcher as those of a theropod rather than a ceratopsid. Henry Fairfield Osborn recognized the similarity between ''Manospondylus gigas'' and ''T. rex'' as early as 1917, by which time the second vertebra had been lost. Owing to the fragmentary nature of the ''Manospondylus'' vertebrae, Osborn did not synonymize the two genera, instead considering the older genus indeterminate. In June 2000, the Black Hills Institute found around 10% of a ''Tyrannosaurus'' skeleton ( BHI 6248) at a site that might have been the original ''M. gigas'' locality.


Skeleton discovery and naming

Barnum Brown Barnum Brown (February 12, 1873 – February 5, 1963), commonly referred to as Mr. Bones, was an American paleontologist. He discovered the first documented remains of ''Tyrannosaurus'' during a career that made him one of the most famous fossil ...
, assistant curator of the
American Museum of Natural History The American Museum of Natural History (AMNH) is a natural history museum on the Upper West Side of Manhattan in New York City. Located in Theodore Roosevelt Park, across the street from Central Park, the museum complex comprises 21 interconn ...
, found the first partial skeleton of ''T. rex'' in eastern Wyoming in 1900. Brown found another partial skeleton in the Hell Creek Formation in Montana in 1902, comprising approximately 34 fossilized bones. Writing at the 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 In ecology, a marsh is a wetland that is dominated by herbaceous plants rather than by woody plants.Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. 497 p More in genera ...
. ... I have never seen anything like it from the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
." Henry Fairfield Osborn, president of the
American Museum of Natural History The American Museum of Natural History (AMNH) is a natural history museum on the Upper West Side of Manhattan in New York City. Located in Theodore Roosevelt Park, across the street from Central Park, the museum complex comprises 21 interconn ...
, named the second skeleton ''T. rex'' in 1905. The generic name is derived from the Greek words (, meaning "tyrant") and (, meaning "lizard"). Osborn used the
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
word , 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. Osborn named the other specimen ''Dynamosaurus imperiosus'' in a paper in 1905. In 1906, Osborn recognized that the two skeletons were from the same species and selected ''Tyrannosaurus'' as the preferred name. In 1941, the ''T. rex'' type specimen was sold to the Carnegie Museum of Natural History in Pittsburgh, Pennsylvania, for $7,000. The original ''Dynamosaurus'' material now resides in the collections of the Natural History Museum, London. ''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. From the 1910s through the end of the 1950s, Barnum's discoveries remained the only specimens of ''Tyrannosaurus'', as the
Great Depression The Great Depression was a severe global economic downturn from 1929 to 1939. The period was characterized by high rates of unemployment and poverty, drastic reductions in industrial production and international trade, and widespread bank and ...
and wars kept many paleontologists out of the field.


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. 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 all previous years, including two of the most complete skeletons found to date: Sue and Stan. Sue Hendrickson, an
amateur An amateur () is generally considered a person who pursues an avocation independent from their source of income. Amateurs and their pursuits are also described as popular, informal, autodidacticism, self-taught, user-generated, do it yourself, DI ...
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. From 1998 to 1999, Field Museum of Natural History staff spent over 25,000 hours taking the rock off the bones. The bones were then shipped to
New Jersey New Jersey is a U.S. state, state located in both the Mid-Atlantic States, Mid-Atlantic and Northeastern United States, Northeastern regions of the United States. Located at the geographic hub of the urban area, heavily urbanized Northeas ...
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. 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. This tyrannosaur also had many bone pathologies, including 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. 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, tall and 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. In the summer of 2000, crews organized by Jack Horner discovered five ''Tyrannosaurus'' skeletons near the Fort Peck Reservoir. 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. Dubbed Jane (BMRP 2002.4.1), the find was thought to be the first known skeleton of a
pygmy In anthropology, pygmy peoples are ethnic groups whose average height is unusually short. The term pygmyism is used to describe the phenotype of endemic short stature (as opposed to disproportionate dwarfism occurring in isolated cases in a po ...
tyrannosaurid, '' Nanotyrannus'', but subsequent research revealed that it is more likely a juvenile ''Tyrannosaurus'', and the most complete juvenile example known; Jane is exhibited at the Burpee Museum of Natural History. 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 In computer technology and telecommunications, online indicates a state of connectivity, and offline indicates a disconnected state. In modern terminology, this usually refers to an Internet connection, but (especially when expressed as "on lin ...
''Tyrannosaurus'' excavation providing daily reports, photos, and video. In 2006, Montana State University revealed that it possessed the largest ''Tyrannosaurus'' skull yet discovered (from a specimen named MOR 008), measuring long. Subsequent comparisons indicated that the longest head was (from specimen LACM 23844) and the widest head was (from Sue).


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 Hadrosaurids (), also hadrosaurs or duck-billed dinosaurs, are members of the ornithischian family Hadrosauridae. This group is known as the duck-billed dinosaurs for the flat duck-bill appearance of the bones in their snouts. The ornithopod fami ...
, 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 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 long by wide. 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 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. A set of footprints in Glenrock, Wyoming dating to the
Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
stage 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 . A follow-up paper appeared in 2017, increasing the speed estimations by 50–80%.


Description


Size

''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 long, was tall at the hips, and according to the most recent studies, using a variety of techniques, maximum body masses have been estimated approximately . A specimen nicknamed Scotty (RSM P2523.8), located at the Royal Saskatchewan Museum, is reported to measure in length. Using a mass estimation technique that extrapolates from the
circumference In geometry, the circumference () is the perimeter of a circle or ellipse. The circumference is the arc length of the circle, as if it were opened up and straightened out to a line segment. More generally, the perimeter is the curve length arou ...
of the femur, Scotty was estimated as the largest known specimen at in body mass. Not every adult ''Tyrannosaurus'' specimen recovered is as big. Historically average adult mass estimates have varied widely over the years, from as low as , to more than , with most modern estimates ranging between . A 2024 study found that there was little evidence of size-based sexual dimorphism in ''T. rex''.


Skull

The largest known ''T. rex'' skulls measure up to in length. 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 Binocular vision is seeing with two eyes. The Field_of_view, field of view that can be surveyed with two eyes is greater than with one eye. To the extent that the visual fields of the two eyes overlap, #Depth, binocular depth can be perceived. Th ...
. The skull bones were massive and the nasals and some other bones were 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. 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. The teeth of ''T. rex'' displayed marked heterodonty (differences in shape). The
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammals h ...
ry 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 bananas" rather than daggers, more widely spaced and also had reinforcing ridges. Those in the upper jaw, twelve per side in mature individuals, were larger than their counterparts of the lower jaw, except at the rear. The largest found so far is estimated to have been long including the root when the animal was alive, making it the largest tooth of any carnivorous dinosaur yet found. The lower jaw was robust. Its front dentary bone bore thirteen teeth. Behind the tooth row, the lower jaw became notably taller. 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 or evidence of extra-oral structures such as scales or potentially lips, with subsequent research on theropod tooth wear patterns supporting such a proposition.


Skeleton

The
vertebral column The spinal column, also known as the vertebral column, spine or backbone, is the core part of the axial skeleton in vertebrates. The vertebral column is the defining and eponymous characteristic of the vertebrate. The spinal column is a segmente ...
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 numbered at least forty. Sue was mounted with forty-seven of such caudal vertebrae. 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 concave rear. The vertebral bodies had single pleurocoels, pneumatic depressions created by air sacs, on their sides. 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, 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. 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 A coracoid is a paired bone which is part of the shoulder assembly in all vertebrates except therian mammals (marsupials and placentals). In therian mammals (including humans), a coracoid process is present as part of the scapula, but this is n ...
, which was rounded. Both shoulder blades were connected by a small furcula. The paired breast bones possibly were made of
cartilage Cartilage is a resilient and smooth type of connective tissue. Semi-transparent and non-porous, it is usually covered by a tough and fibrous membrane called perichondrium. In tetrapods, it covers and protects the ends of long bones at the joints ...
only. 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 The ulna or ulnar bone (: ulnae or ulnas) is a long bone in the forearm stretching from the elbow to the wrist. It is on the same side of the forearm as the little finger, running parallel to the Radius (bone), radius, the forearm's other long ...
and radius, were straight elements, much shorter than the humerus. The second
metacarpal In human anatomy, the metacarpal bones or metacarpus, also known as the "palm bones", are the appendicular bones that form the intermediate part of the hand between the phalanges (fingers) and the carpal bones ( wrist bones), which articulate ...
was longer and wider than the first, whereas normally in theropods the opposite is true. The forelimbs had only two clawed fingers, along with an additional splint-like small third
metacarpal In human anatomy, the metacarpal bones or metacarpus, also known as the "palm bones", are the appendicular bones that form the intermediate part of the hand between the phalanges (fingers) and the carpal bones ( wrist bones), which articulate ...
representing the remnant of a third digit. The
pelvis The pelvis (: pelves or pelvises) is the lower part of an Anatomy, anatomical Trunk (anatomy), trunk, between the human abdomen, abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also c ...
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 The ischium (; : is ...
was slender and straight, pointing obliquely to behind and below. In contrast to the arms, the hindlimbs were among the longest in proportion to body size of any theropod. In the 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. Compensating for the immense bulk of the animal, many bones throughout the skeleton were hollowed, reducing its weight without significant loss of strength.


Classification

''Tyrannosaurus'' is the type genus of the superfamily Tyrannosauroidea, the
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
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
Asia Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...
n '' Tarbosaurus'', both of which have occasionally been synonymized with ''Tyrannosaurus''. Tyrannosaurids were once commonly thought to be descendants of earlier large theropods such as megalosaurs and carnosaurs, although more recently they were reclassified with the generally smaller coelurosaurs. 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 The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
. 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. 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. Many
phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
analyses have found '' Tarbosaurus bataar'' to be the sister taxon of ''T. rex''. 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. 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''. Below is the cladogram of Tyrannosauridae based on the phylogenetic analysis conducted by Loewen and colleagues in 2013. 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 In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
clade containing ''Daspletosaurus'' and '' Thanatotheristes'', typically referred to as the Daspletosaurini.


Additional species

In 1955, Soviet
paleontologist Paleontology, also spelled as palaeontology or palæontology, is the scientific study of the life of the past, mainly but not exclusively through the study of fossils. Paleontologists use fossils as a means to classify organisms, measure geolo ...
Evgeny Maleev named a new species, ''Tyrannosaurus bataar'', from
Mongolia Mongolia is a landlocked country in East Asia, bordered by Russia to the north and China to the south and southeast. It covers an area of , with a population of 3.5 million, making it the world's List of countries and dependencies by po ...
. By 1965, this species was renamed as a distinct genus, ''Tarbosaurus bataar''. 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 are similar enough to be considered members of the same genus, restoring the Mongolian
taxon In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
's original binomial name. Some specimens from the Late Cretaceous deposits of
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
have been described as new species of ''Tyrannosaurus'': ''T. lanpingensis'' based on isolated
lateral Lateral is a geometric term of location which may also refer to: Biology and healthcare * Lateral (anatomy), a term of location meaning "towards the side" * Lateral cricoarytenoid muscle, an intrinsic muscle of the larynx * Lateral release ( ...
tooth from the red beds of
Yunnan Yunnan; is an inland Provinces of China, province in Southwestern China. The province spans approximately and has a population of 47.2 million (as of 2020). The capital of the province is Kunming. The province borders the Chinese provinces ...
in 1975; ''T. turpanensis'' from the Subashi Formation, Turpan Basin,
Xinjiang Xinjiang,; , SASM/GNC romanization, SASM/GNC: Chinese postal romanization, previously romanized as Sinkiang, officially the Xinjiang Uygur Autonomous Region (XUAR), is an Autonomous regions of China, autonomous region of the China, People' ...
in 1978; and ''T. luanchuanensis'' from the Quiba Formation, Tantou Basin,
Henan Province Henan; alternatively Honan is a province in Central China. Henan is home to many heritage sites, including Yinxu, the ruins of the final capital of the Shang dynasty () and the Shaolin Temple. Four of the historical capitals of China, Luo ...
in 1979–1980. All these taxa were published without detailed descriptions and were later accepted as junior synonyms of ''Tarbosaurus bataar'' by Holtz in 2004. VGI, no. 231/3, a large
phalanx bone The phalanges (: phalanx ) are digital bones in the hands and feet of most vertebrates. In primates, the thumbs and big toes have two phalanges while the other digits have three phalanges. The phalanges are classed as long bones. Structu ...
, 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 In human anatomy, the metacarpal bones or metacarpus, also known as the "palm bones", are the appendicular bones that form the intermediate part of the hand between the phalanges (fingers) and the carpal bones ( wrist bones), which articulate ...
I or metatarsal I that possibly belongs to ceratosaur. In their 2023 overview, Averianov and Lopatin mention this specimen as well as a single tooth from the same site only as Theropoda indet. 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 A synonym is a word, morpheme, or phrase that means precisely or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are a ...
with ''Zhuchengtyrannus''. In any case, ''T. zhuchengensis'' is considered to be a ''
nomen dubium In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application. Zoology In case of a ''nomen dubium,'' it may be impossible to determine whether a ...
'' as the holotype lacks
diagnostic Diagnosis (: diagnoses) is the identification of the nature and cause of a certain phenomenon. Diagnosis is used in a lot of different academic discipline, disciplines, with variations in the use of logic, analytics, and experience, to determine " ...
features below the level Tyrannosaurinae. 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. In 2018, a paper describing tyrannosaurid teeth from the Two Medicine Formation noted a premaxillary tooth (YPM VPPU 023469) had a strong resemblance to the teeth of Sue to the exclusion of any Campanian tyrannosaurid. Additionally, the authors of this paper suggested that CM 9401 also comes from the Two Medicine Formation, as there were preservational similarities between its locality and the Willow Creek anticline, which is where the tooth was found. Notably, this would place both specimens in the Flag Butte Member of the Two Medicine Formation, which dates from 77 to 76.3 Ma, far younger than any other ''Tyrannosaurus'' specimen, and directly contemporaneous with ''Daspletosaurus.'' In 2025, these specimens, with their old geologic age, were used as evidence by Charlie Scherer to suggest that the Tyrannosaurini did not evolve directly from ''Daspletosaurus''. 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 The femur (; : femurs or femora ), or thigh bone is the only long bone, bone in the thigh — the region of the lower limb between the hip and the knee. In many quadrupeds, four-legged animals the femur is the upper bone of the hindleg. The Femo ...
(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''. 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. Their criticism was subsequently published in a technical paper. 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. Paul still rejected the objections raised by critics, insisting that they are unwilling to consider that ''Tyrannosaurus'' might represent more than one species.Osborne M
"Study Refutes Controversial Research That Divided the T. Rex Into Three Species"
. Smithsonian Magazine July 27, 2022


''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 A palatine or palatinus (Latin; : ''palatini''; cf. derivative spellings below) is a high-level official attached to imperial or royal courts in Europe since Roman Empire, Roman times.
, and an incomplete
maxilla In vertebrates, the maxilla (: maxillae ) is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The two maxil ...
from the skull, the left
dentary In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone ...
, right
splenial The splenial is a small bone in the lower jaw of reptile Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology ...
, right prearticular, right angular and right articular from the lower jaws, isolated teeth, and chevrons. Some of the bones were briefly mentioned in 1984 as belonging to ''T. rex'', and described in 1986. Lehman and Carpenter (1990) suggested that NMMNH P-3698 belonged to a new tyrannosaurid genus, while Carr and Williamson (2000) disagreed with their claim. 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 probable association with the '' Alamosaurus'' fossil. 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. 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. The rock layers were initially estimated to date to between 72.7 and 70.9 Ma, correlating to the latest Campanian or earliest Maastrichtian. However, in a 2024
conference A conference is a meeting, often lasting a few days, which is organized on a particular subject, or to bring together people who have a common interest. Conferences can be used as a form of group decision-making, although discussion, not always d ...
abstracts, the specific stratigraphic layer which yielded ''T. mcraeensis'' was estimated to be younger than 69.0 ± 0.4 Ma and older than 66.0 Ma based on the sandstone from the same fossil locality, which would suggest that the age of ''T. mcraeensis'' is much closer to ''T. rex'' than previously thought. ''T. mcraeensis'' was estimated at 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''. In 2025, Morrison and colleagues noted the uncertainty in the previous age estimate of ''T. mcraeensis''. The authors argued that the inclusion of the titanosaurian fossil (cf. ''Alamosaurus'') as a biostratigraphic marker of the Hall Lake Formation fauna makes the dating problematic due to the fossil having a huge variability in assumed sedimentation rates. The study also suggested the uncertainty in the morphology of ''T. mcraeensis'' holotype being reliably outside the known variation of ''T. rex''. Regardless of the age and validity of ''T. mcraeensis'', the authors concluded that the genus ''Tyrannosaurus'' still most likely originated from Laramidia and that the ancestors of this genus possibly migrated from Asia. Later that year, Voris and colleagues regarded ''T. mcraeensis'' as a
junior synonym In taxonomy, the scientific classification of living organisms, a synonym is an alternative scientific name for the accepted scientific name of a taxon. The botanical and zoological codes of nomenclature treat the concept of synonymy differently. ...
of ''T. rex'', agreeing with the problematic stratigraphic data and finding all of the proposed diagnostic characters of ''T. mcraeensis'' to be in the known ''T. rex'' specimen range of variation.


''Nanotyrannus''

Other tyrannosaurid fossils found in the same formations as ''T. rex'' were originally classified as separate taxa, including ''Aublysodon'' and ''Albertosaurus megagracilis'', the latter being named ''Dinotyrannus megagracilis'' in 1995. These fossils are now universally considered to belong to juvenile ''T. rex''. A small but nearly complete skull from Montana, 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. 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 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 long when it died. However, In 1999, a detailed analysis by Thomas Carr revealed the specimen to be a juvenile, leading Carr and many other paleontologists to consider it a juvenile ''T. rex'' individual. 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 discovery of "Jane" as a confirmation that ''Nanotyrannus'' was, in fact, a juvenile ''T. rex''.Currie, Henderson, Horner and Williams (2005). "On tyrannosaur teeth, tooth positions and the taxonomic status of ''Nanotyrannus lancensis''." In "The origin, systematics, and paleobiology of Tyrannosauridae", a symposium hosted jointly by Burpee Museum of Natural History and Northern Illinois University.Henderson (2005). "Nano No More: The death of the pygmy tyrant." In "The origin, systematics, and paleobiology of Tyrannosauridae", a symposium hosted jointly by Burpee Museum of Natural History and Northern Illinois University. Peter Larson continued to support the hypothesis that ''N''. ''lancensis'' was a separate but a 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.Larson (2005). "A case for ''Nanotyrannus''." In "The origin, systematics, and paleobiology of Tyrannosauridae", a symposium hosted jointly by Burpee Museum of Natural History and Northern Illinois University.Larson P (2013), "The validity of Nanotyrannus Lancensis (Theropoda, Lancian – Upper Maastrichtian of North America)", Society of Vertebrate Paleontology: 73rd annual meeting, ''Abstracts with Programs'', p. 159. Later research revealed that other tyrannosaurids such as ''Gorgosaurus'' also experienced reduction in tooth count during growth, 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. 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. In 2016, the analysis of limb proportions by Persons and Currie suggested ''Nanotyrannus'' specimens to have differing cursoriality levels, potentially separating it from ''T. rex''. However, paleontologist Manabu Sakomoto has commented that this conclusion may be impacted by low
sample size Sample size determination or estimation is the act of choosing the number of observations or replicates to include in a statistical sample. The sample size is an important feature of any empirical study in which the goal is to make inferences abo ...
, and the discrepancy does not necessarily reflect taxonomic distinction. 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. The same year, Carr and colleagues noted that this was insufficient to clarify ''Nanotyrannus''' validity or classification, being a common and ontogenetically variable feature among 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''. The same year, Carr published a paper on ''T''. ''rex'''s growth history, finding that 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 . 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. 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." In 2024, Nick Longrich and Evan Thomas Saitta re-examined 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. Voris et al. (2025) suggested that the scoring of ''Nanotyrannus'' in a more basal position by Longrich & Saitta (2024) would have been caused by "scoring characteristics related to immaturity" and problematic data sets, supporting the previous interpretation that ''Nanotyrannus'' represents a juvenile ''T. rex''.Supplementary Information
/ref>


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 , while the largest adults, such as FMNH PR2081 (Sue) most likely weighed about . Histologic analysis of ''T. rex'' bones showed LACM 28471 had aged only 2 years when it died. Sue was initially estimated at 28 years old, an age which was at the time considered close to the maximum for the species, but the most recent studies estimate Sue's age at 27-33 years. Similarly, Trix (RGM 792.000) is estimated to have been at least 30 years old at time of death. 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 with their mass. A ''T. rex'' growth curve is S-shaped, with juveniles remaining under until approximately 14 years of age, when body size began to increase dramatically. During this rapid growth phase, a young ''T. rex'' would gain an average of a year for the next four years. At 18 years of age, the curve plateaus again, indicating that growth slowed dramatically. For example, only separated the 28-year-old Sue from a 22-year-old Canadian specimen ( RTMP 81.12.1). A 2004 histological study performed by different workers corroborates these results, finding that rapid growth began to slow at around 16 years of age. 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". 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. 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 The femur (; : femurs or femora ), or thigh bone is the only long bone, bone in the thigh — the region of the lower limb between the hip and the knee. In many quadrupeds, four-legged animals the femur is the upper bone of the hindleg. The Femo ...
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. Further study indicates an age of 18 for this specimen. 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. Other tyrannosaurids exhibit extremely similar growth curves, although with lower growth rates corresponding to their lower adult sizes. 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 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. 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 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 A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
or to the bias of fossil collectors towards larger, more spectacular specimens. 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. Gregory S. Paul also writes that ''Tyrannosaurus'' reproduced quickly and died young but attributes their short lifespans to the dangerous lives they lived.


Skin and possible filamentous feathering

The discovery of feathered dinosaurs led to debate regarding whether, and to what extent, ''Tyrannosaurus'' might have been feathered. Filamentous structures, which are commonly recognized as the precursors of
feather Feathers are epidermal growths that form a distinctive outer covering, or plumage, on both avian (bird) and some non-avian dinosaurs and other archosaurs. They are the most complex integumentary structures found in vertebrates and an exa ...
s, have been reported in the small-bodied, basal tyrannosauroid '' Dilong paradoxus'' from the Early Cretaceous Yixian Formation of China in 2004. Because
integument In biology, an integument is the tissue surrounding an organism's body or an organ within, such as skin, a husk, Exoskeleton, shell, germ or Peel (fruit), rind. Etymology The term is derived from ''integumentum'', which is Latin for "a coverin ...
ary impressions of larger tyrannosauroids known at that time showed evidence of
scales Scale or scales may refer to: Mathematics * Scale (descriptive set theory), an object defined on a set of points * Scale (ratio), the ratio of a linear dimension of a model to the corresponding dimension of the original * Scale factor, a number ...
, the researchers who studied ''Dilong'' speculated that insulating feathers might have been lost by larger species due to their smaller surface-to-volume ratio. The subsequent discovery of the giant species '' 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. 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. The study concluded that feather covering of large tyrannosaurids such as ''Tyrannosaurus'' was, if present, limited to the upper side of the trunk. 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
crocodilia Crocodilia () is an order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorph pseudosuchia ...
ns. 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. 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. 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. 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
squamate Squamata (, Latin ''squamatus'', 'scaly, having scales') is the largest Order (biology), order of reptiles; most members of which are commonly known as Lizard, lizards, with the group also including Snake, snakes. With over 11,991 species, it i ...
s such as monitor lizards or marine iguanas than those of modern
crocodilia Crocodilia () is an order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorph pseudosuchia ...
ns like
alligator An alligator, or colloquially gator, is a large reptile in the genus ''Alligator'' of the Family (biology), family Alligatoridae in the Order (biology), order Crocodilia. The two Extant taxon, extant species are the American alligator (''A. mis ...
s. 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.


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 Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
in ''T. rex'', with the 'robust' morph usually suggested to be female. For example, the
pelvis The pelvis (: pelves or pelvises) is the lower part of an Anatomy, anatomical Trunk (anatomy), trunk, between the human abdomen, abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also c ...
of several 'robust' specimens seemed to be wider, perhaps to allow the passage of eggs. It was also thought 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
crocodile Crocodiles (family (biology), family Crocodylidae) or true crocodiles are large, semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term "crocodile" is sometimes used more loosely to include ...
s. 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. 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 Saskatchewan is a Provinces and territories of Canada, province in Western Canada. It is bordered on the west by Alberta, on the north by the Northwest Territories, on the east by Manitoba, to the northeast by Nunavut, and to the south by the ...
to
New Mexico New Mexico is a state in the Southwestern United States, Southwestern region of the United States. It is one of the Mountain States of the southern Rocky Mountains, sharing the Four Corners region with Utah, Colorado, and Arizona. It also ...
, 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. Only a single ''Tyrannosaurus'' specimen has been conclusively shown to belong to a specific sex. Examination of B-rex demonstrated the preservation of
soft tissue Soft tissue connective tissue, connects and surrounds or supports internal organs and bones, and includes muscle, tendons, ligaments, Adipose tissue, fat, fibrous tissue, Lymphatic vessel, lymph and blood vessels, fasciae, and synovial membranes. ...
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 Calcium is a chemical element; it has symbol Ca and atomic number 20. As an alkaline earth metal, calcium is a reactive metal that forms a dark oxide-nitride layer when exposed to air. Its physical and chemical properties are most similar to it ...
for the production of
eggshell An eggshell is the outer covering of a hard-shelled egg (biology), egg and of some forms of eggs with soft outer coats. Worm eggs Nematode eggs present a two layered structure: an external vitellin layer made of chitin that confers mechanical ...
during
ovulation Ovulation is an important part of the menstrual cycle in female vertebrates where the egg cells are released from the ovaries as part of the ovarian cycle. In female humans ovulation typically occurs near the midpoint in the menstrual cycle and ...
. 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
hormone A hormone (from the Ancient Greek, Greek participle , "setting in motion") is a class of cell signaling, signaling molecules in multicellular organisms that are sent to distant organs or tissues by complex biological processes to regulate physio ...
s like
estrogen Estrogen (also spelled oestrogen in British English; see spelling differences) is a category of sex hormone responsible for the development and regulation of the female reproductive system and secondary sex characteristics. There are three ...
. This strongly suggests that B-rex was female and that she died during ovulation. 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
evolution Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
ary relationship between the two.


Posture

Like many
bipedal Bipedalism is a form of terrestrial locomotion where an animal moves by means of its two rear (or lower) limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' ...
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 Kangaroos are marsupials from the family Macropodidae (macropods, meaning "large foot"). In common use, the term is used to describe the largest species from this family, the red kangaroo, as well as the antilopine kangaroo, eastern gre ...
. This concept dates from Joseph Leidy's 1865 reconstruction of '' Hadrosaurus'', the first to depict a dinosaur in a bipedal posture. 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. 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 In materials science, a dislocation or Taylor's dislocation is a linear crystallographic defect or irregularity within a crystal structure that contains an abrupt change in the arrangement of atoms. The movement of dislocations allow atoms to sli ...
or weakening of several
joint A joint or articulation (or articular surface) is the connection made between bones, ossicles, or other hard structures in the body which link an animal's skeletal system into a functional whole.Saladin, Ken. Anatomy & Physiology. 7th ed. McGraw- ...
s, including the hips and the articulation between the head and the
spinal column The spinal column, also known as the vertebral column, spine or backbone, is the core part of the axial skeleton in vertebrates. The vertebral column is the defining and eponymous characteristic of the vertebrate. The spinal column is a segmen ...
. 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 Yale University is a Private university, private Ivy League research university in New Haven, Connecticut, United States. Founded in 1701, Yale is the List of Colonial Colleges, third-oldest institution of higher education in the United Stat ...
's Peabody Museum of Natural History) until the 1990s, when films such as ''
Jurassic Park ''Jurassic Park'', later referred to as ''Jurassic World'', is an American science fiction media franchise created by Michael Crichton, centered on a disastrous attempt to create a theme park of De-extinction#Cloning, cloned dinosaurs. It bega ...
'' introduced a more accurate posture to the general public. 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. 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. 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 and as evidence that they were indeed used and acquired wounds,Stevens K.A., Larson P, Willis E.D. & Anderson A. "Rex, sit: digital modeling of ''Tyrannosaurus rex'' at rest". In Larson P & Carpenter K (eds.). ''Tyrannosaurus rex, the tyrant king'' (Indiana University Press, 2008). p. 192-203 like the rest of the body.


Arms

When ''T. rex'' was first discovered, the
humerus The humerus (; : humeri) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius (bone), radius and ulna, and consists of three sections. The humeral upper extrem ...
was the only element of the forelimb known. Retrieved October 6, 2008. For the initial mounted skeleton as seen by the public in 1915, Osborn substituted longer, three-fingered forelimbs like those of '' Allosaurus''. A year earlier, Lawrence Lambe described the short, two-fingered forelimbs of the closely related ''Gorgosaurus''. This strongly suggested that ''T. rex'' had similar forelimbs, but this
hypothesis A hypothesis (: hypotheses) is a proposed explanation for a phenomenon. A scientific hypothesis must be based on observations and make a testable and reproducible prediction about reality, in a process beginning with an educated guess o ...
was not confirmed until the first complete ''T. rex'' forelimbs were identified in 1989, belonging to MOR 555 (the "Wankel rex"). The remains of Sue also include complete forelimbs. ''T. rex'' arms are very small relative to overall body size, measuring only long, and some scholars have labelled them as vestigial. However, the bones show large areas for
muscle Muscle is a soft tissue, one of the four basic types of animal tissue. There are three types of muscle tissue in vertebrates: skeletal muscle, cardiac muscle, and smooth muscle. Muscle tissue gives skeletal muscles the ability to muscle contra ...
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. Newman (1970) suggested that the forelimbs were used to assist ''Tyrannosaurus'' in rising from a prone position. Since then, other functions have been proposed, although some scholars find them implausible. 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.Padian K (2022)
"Why tyrannosaurid forelimbs were so short: An integrative hypothesis"
. ''Acta Palaeontologica Polonica'' 67(1): p. 63-76
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 A bone is a rigid organ that constitutes part of the skeleton in most vertebrate animals. Bones protect the various other organs of the body, produce red and white blood cells, store minerals, provide structure and support for the body, an ...
, 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 by itself; other muscles such as the brachialis would work along with the biceps to make elbow 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 ''Deinonychus'' ( ; ) is a genus of Dromaeosauridae, dromaeosaurid Theropoda, theropod dinosaur with one described species, ''Deinonychus antirrhopus''. This species, which could grow up to long, lived during the early Cretaceous Period (ge ...
'' 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. 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. This was dismissed by Padian, who argued that Stanley based his conclusion on incorrectly estimated forelimb size and range of motion.


Thermoregulation

''Tyrannosaurus'', like most dinosaurs, was long thought to have an
ectotherm An ectotherm (), more commonly referred to as a "cold-blooded animal", is an animal in which internal physiological sources of heat, such as blood, are of relatively small or of quite negligible importance in controlling body temperature.Dav ...
ic ("cold-blooded") reptilian
metabolism Metabolism (, from ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run cellular processes; the co ...
. The idea of dinosaur ectothermy was challenged by scientists like Robert T. Bakker and John Ostrom in the early years of the " Dinosaur Renaissance", beginning in the late 1960s. ''T. rex'' itself was claimed to have been
endothermic An endothermic process is a chemical or physical process that absorbs heat from its surroundings. In terms of thermodynamics, it is a thermodynamic process with an increase in the enthalpy (or internal energy ) of the system.Oxtoby, D. W; Gillis, ...
("warm-blooded"), implying a very active lifestyle. Since then, several paleontologists have sought to determine the ability of ''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
vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
s. 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 between the vertebrae of the torso and the
tibia The tibia (; : tibiae or tibias), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two Leg bones, bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outsi ...
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. 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). 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'').
Ornithischia Ornithischia () is an extinct clade of mainly herbivorous dinosaurs characterized by a pelvic structure superficially similar to that of birds. The name ''Ornithischia'', or "bird-hipped", reflects this similarity and is derived from the Greek ...
n dinosaurs also showed evidence of homeothermy, while varanid
lizard Lizard is the common name used for all Squamata, squamate reptiles other than snakes (and to a lesser extent amphisbaenians), encompassing over 7,000 species, ranging across all continents except Antarctica, as well as most Island#Oceanic isla ...
s from the same formation did not. In 2022, Wiemann and colleagues used a different approach—the
spectroscopy Spectroscopy is the field of study that measures and interprets electromagnetic spectra. In narrower contexts, spectroscopy is the precise study of color as generalized from visible light to all bands of the electromagnetic spectrum. Spectro ...
of lipoxidation signals, which are byproducts of
oxidative phosphorylation Oxidative phosphorylation(UK , US : or electron transport-linked phosphorylation or terminal oxidation, is the metabolic pathway in which Cell (biology), cells use enzymes to Redox, oxidize nutrients, thereby releasing chemical energy in order ...
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. 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 turtle Sea turtles (superfamily Chelonioidea), sometimes called marine turtles, are reptiles of the order Testudines and of the suborder Cryptodira. The seven existing species of sea turtles are the flatback, green, hawksbill, leatherback, loggerh ...
s. Similar to contemporary crocodilians, openings (dorsotemporal fenestrae) in the skull roofs of ''Tyrannosaurus'' may have aided thermoregulation.


Soft tissue

In the March 2005 issue of ''
Science Science is a systematic discipline that builds and organises knowledge in the form of testable hypotheses and predictions about the universe. Modern science is typically divided into twoor threemajor branches: the natural sciences, which stu ...
'', Mary Higby Schweitzer of
North Carolina State University North Carolina State University (NC State, North Carolina State, NC State University, or NCSU) is a public university, public Land-grant university, land-grant research university in Raleigh, North Carolina, United States. Founded in 1887 and p ...
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. 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 vessel Blood vessels are the tubular structures of a circulatory system that transport blood throughout many Animal, animals’ bodies. Blood vessels transport blood cells, nutrients, and oxygen to most of the Tissue (biology), tissues of a Body (bi ...
s and fibrous but elastic
bone A bone is a rigid organ that constitutes part of the skeleton in most vertebrate animals. Bones protect the various other organs of the body, produce red and white blood cells, store minerals, provide structure and support for the body, ...
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, 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. 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. Research on some of the tissues involved has suggested that birds are closer relatives to tyrannosaurs than other modern animals. 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. In studies reported in ''Science'' in April 2007, Asara and colleagues concluded that seven traces of
collagen Collagen () is the main structural protein in the extracellular matrix of the connective tissues of many animals. It is the most abundant protein in mammals, making up 25% to 35% of protein content. Amino acids are bound together to form a trip ...
proteins detected in purified ''T. rex'' bone most closely match those reported in
chicken The chicken (''Gallus gallus domesticus'') is a domesticated subspecies of the red junglefowl (''Gallus gallus''), originally native to Southeast Asia. It was first domesticated around 8,000 years ago and is now one of the most common and w ...
s, 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". The presumed soft tissue was called into question by Thomas Kaye of the
University of Washington The University of Washington (UW and informally U-Dub or U Dub) is a public research university in Seattle, Washington, United States. Founded in 1861, the University of Washington is one of the oldest universities on the West Coast of the Uni ...
and his co-authors in 2008. They contend that what was really inside the tyrannosaur bone was slimy
biofilm A biofilm is a Syntrophy, syntrophic Microbial consortium, community of microorganisms in which cell (biology), cells cell adhesion, stick to each other and often also to a surface. These adherent cells become embedded within a slimy ext ...
created by bacteria that coated the voids once occupied by blood vessels and cells. The researchers found that what previously had been identified as remnants of blood cells, because of the presence of iron, were actually
framboid A framboid is a micromorphological feature common to certain sedimentary minerals, particularly pyrite (FeS2). The first known use of the term is ascribed to Rust in 1935 and is derived from the French 'framboise', meaning 'raspberry', reflecting th ...
s, microscopic mineral spheres bearing iron. They found similar spheres in a variety of other fossils from various periods, including an
ammonite Ammonoids are extinct, (typically) coiled-shelled cephalopods comprising the subclass Ammonoidea. They are more closely related to living octopuses, squid, and cuttlefish (which comprise the clade Coleoidea) than they are to nautiluses (family N ...
. 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. 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. 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. Other research challenges the identification of soft tissue as biofilm and confirms finding "branching, vessel-like structures" from within fossilized bone.


Speed

Scientists have produced a wide range of possible maximum running speeds for ''Tyrannosaurus'': mostly around , but as low as and as high as , though it running this speed is very unlikely. ''Tyrannosaurus'' was a bulky and heavy carnivore so it is unlikely to run very fast at all compared to other theropods like '' Carnotaurus'' or '' Giganotosaurus''. Researchers have relied on various estimating techniques because, while there are many tracks of large theropods walking, none showed evidence of running. A 2002 report used a mathematical model (validated by applying it to three living animals:
alligator An alligator, or colloquially gator, is a large reptile in the genus ''Alligator'' of the Family (biology), family Alligatoridae in the Order (biology), order Crocodilia. The two Extant taxon, extant species are the American alligator (''A. mis ...
s,
chicken The chicken (''Gallus gallus domesticus'') is a domesticated subspecies of the red junglefowl (''Gallus gallus''), originally native to Southeast Asia. It was first domesticated around 8,000 years ago and is now one of the most common and w ...
s, and
human Humans (''Homo sapiens'') or modern humans are the most common and widespread species of primate, and the last surviving species of the genus ''Homo''. They are Hominidae, great apes characterized by their Prehistory of nakedness and clothing ...
s; and eight more species, including emus and ostriches) to gauge the leg muscle mass needed for fast running (over ). 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 or so, or that other animals like ostriches and
horse The horse (''Equus ferus caballus'') is a domesticated, one-toed, hoofed mammal. It belongs to the taxonomic family Equidae and is one of two extant subspecies of ''Equus ferus''. The horse has evolved over the past 45 to 55 mi ...
s with long, flexible legs are able to achieve high speeds through slower but longer strides. Proposed top speeds exceeded 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 for walking or jogging would have been possible. 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). 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. Together, these leg features allowed ''Tyrannosaurus'' to transmit locomotory forces from the foot to the lower leg more effectively than in earlier theropods. 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 , top running speed is limited by body size, so longer legs instead were found to have correlated with 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 reduced energy expenditures during hunting or scavenging. This in turn likely resulted in tyrannosaurs having a reduced 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. A study published in 2021 by Pasha van Bijlert et al., calculated the preferred walking speed of ''Tyrannosaurus'', reporting a speed of . 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 Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
, 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. A 2017 study estimated the top running speed of ''Tyrannosaurus'' as , speculating that ''Tyrannosaurus'' exhausted its energy reserves long before reaching top speed, resulting in a parabola-like relationship between size and speed. 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 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''. 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 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.


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 2021, show a large tyrannosaurid rising from a prone position by rising up using its elbows in conjunction with the pads on their feet to stand. These two unique sets of fossils were found in Ludlow, Colorado and Cimarron, New Mexico. 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 , significantly faster than previously assumed for estimations of walking speed in tyrannosaurids.


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. 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 acuity of a human and surpassed the visual acuity of an eagle, which is 3.6 times that of a person. Stevens estimated a limiting far point (that is, the distance at which an object can be seen as separate from the horizon) as far as away, which is greater than the that a human can see. 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 ''Triceratops'' ( ; ) is a genus of Chasmosaurinae, chasmosaurine Ceratopsia, ceratopsian dinosaur that lived during the late Maastrichtian age of the Late Cretaceous Period (geology), period, about 68 to 66 million years ago on the island ...
'', 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''. 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 ~ in diameter with an internal aperture diameter of ~. Based on eye proportions in living reptiles, this implies a pupil diameter of about , an iris diameter about that of the sclerotic ring, and an eyeball diameter of . Carpenter also estimated an eyeball depth of ~. Based on these calculations, the
f-number An f-number is a measure of the light-gathering ability of an optical system such as a camera lens. It is calculated by dividing the system's focal length by the diameter of the entrance pupil ("clear aperture").Smith, Warren ''Modern Optical ...
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. ''Tyrannosaurus'' had very large
olfactory bulb The olfactory bulb (Latin: ''bulbus olfactorius'') is a neural structure of the vertebrate forebrain involved in olfaction, the sense of smell. It sends olfactory information to be further processed in the amygdala, the orbitofrontal cortex (OF ...
s and
olfactory nerve The olfactory nerve, also known as the first cranial nerve, cranial nerve I, or simply CN I, is a cranial nerve that contains sensory nerve fibers relating to the sense of smell. The afferent nerve fibers of the olfactory receptor neurons t ...
s 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. Somewhat unusually among theropods, ''T. rex'' had a very long
cochlea The cochlea is the part of the inner ear involved in hearing. It is a spiral-shaped cavity in the bony labyrinth, in humans making 2.75 turns around its axis, the modiolus (cochlea), modiolus. A core component of the cochlea is the organ of Cort ...
. 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 behavior. A 2017 study by Thomas Carr and colleagues 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 neuron Sensory neurons, also known as afferent neurons, are neurons in the nervous system, that convert a specific type of stimulus, via their receptors, into action potentials or graded receptor potentials. This process is called sensory transduc ...
s. 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. 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 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. 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 ''Spinosaurus'' (; ) is a genus of large spinosaurid theropod dinosaurs that lived in what now is North Africa during the Cenomanian faunal stage, stage of the Late Cretaceous Period (geology), period, about 100 to 94 annum, million year ...
'', '' 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. A study by Grant R. Hurlburt, Ryan C. Ridgely and Lawrence Witmer obtained estimates for
Encephalization Quotient Encephalization quotient (EQ), encephalization level (EL), or just encephalization is a relative brain size measure that is defined as the ratio between observed and predicted brain mass for an animal of a given size, based on nonlinear regre ...
s (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 ''Tyrannosauruss 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. 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 The chimpanzee (; ''Pan troglodytes''), also simply known as the chimp, is a species of Hominidae, great ape native to the forests and savannahs of tropical Africa. It has four confirmed subspecies and a fifth proposed one. When its close rel ...
(2.2–2.5), though this may be debatable as reptilian and mammalian encephalization quotients are not equivalent.


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. A find in
South Dakota South Dakota (; Sioux language, Sioux: , ) is a U.S. state, state in the West North Central states, North Central region of the United States. It is also part of the Great Plains. South Dakota is named after the Dakota people, Dakota Sioux ...
where three ''T. rex'' skeletons were in close proximity may suggest the formation of a pack. Cooperative pack hunting may have been an effective strategy for subduing prey with advanced
anti-predator adaptation Anti-predator adaptations are mechanisms developed through evolution that assist Predation, prey organisms in their constant struggle against predators. Throughout the animal kingdom, adaptations have evolved for every stage of this struggle, na ...
s which pose potential lethality such as ''
Triceratops ''Triceratops'' ( ; ) is a genus of Chasmosaurinae, chasmosaurine Ceratopsia, ceratopsian dinosaur that lived during the late Maastrichtian age of the Late Cretaceous Period (geology), period, about 68 to 66 million years ago on the island ...
'' and '' Ankylosaurus''. Currie's pack-hunting ''T. rex'' hypothesis has been criticized for not having been
peer-review Peer review is the evaluation of work by one or more people with similar competencies as the producers of the work ( peers). It functions as a form of self-regulation by qualified members of a profession within the relevant field. Peer review ...
ed, but rather was discussed in a television interview and book called ''Dino Gangs''. 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 interpreted with reference to evidence in other closely related species. According to Currie gregariousness in ''Albertosaurus sarcophagus'' is supported by the discovery of 26 individuals with varied ages in the Dry Island bonebed. He ruled out the possibility of a predator trap due to the similar preservation state of individuals and the near absence of herbivores. (not printed until 2000) Additional support of tyrannosaurid gregariousness can be found in fossilized trackways from the Upper Cretaceous Wapiti Formation of northeastern
British Columbia British Columbia is the westernmost Provinces and territories of Canada, province of Canada. Situated in the Pacific Northwest between the Pacific Ocean and the Rocky Mountains, the province has a diverse geography, with rugged landscapes that ...
, Canada, left by three tyrannosaurids traveling in the same direction. 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). Others researchers have speculated that instead of large theropod social groups, some of these finds represent behavior more akin to
Komodo dragon The Komodo dragon (''Varanus komodoensis''), also known as the Komodo monitor, is a large reptile of the monitor lizard family Varanidae that is endemic to the Indonesian islands of Komodo (island), Komodo, Rinca, Flores, Gili Dasami, and Gili ...
-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. 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. 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. 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 300 px, Bone tissue is removed by osteoclasts, and then new bone tissue is formed by osteoblasts. Both processes utilize cytokine ( Insulin-like_growth_factor.html" ;"title="TGF-β, Insulin-like growth factor">IGF) signalling. In osteology, bone ...
(not regrowth).


Feeding strategies

Most paleontologists accept that ''Tyrannosaurus'' was both an active
predator Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
and a
scavenger Scavengers are animals that consume Corpse decomposition, dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a he ...
like most large
carnivores A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose nutrition and energy requirements are met by consumption of animal tissues (mainly mu ...
. By far the largest carnivore in its environment, ''T. rex'' was most likely an
apex predator An apex predator, also known as a top predator or superpredator, is a predator at the top of a food chain, without natural predators of its own. Apex predators are usually defined in terms of trophic dynamics, meaning that they occupy the hig ...
, preying upon hadrosaurs, armored herbivores like
ceratopsia Ceratopsia or Ceratopia ( or ; Ancient Greek, Greek: "horned faces") is a group of herbivore, herbivorous, beaked dinosaurs that thrived in what are now North America, Asia and Europe, during the Cretaceous Period (geology), Period, although ance ...
ns and ankylosaurs, and possibly
sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their b ...
s. 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. 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. Even higher estimates were made by Mason B. Meers in 2003. This allowed it to crush bones during repetitive biting and fully consume the carcasses of large dinosaurs. 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. A debate exists, however, about whether ''Tyrannosaurus'' was primarily a
predator Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
or a pure
scavenger Scavengers are animals that consume Corpse decomposition, dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a he ...
. The debate originated in a 1917 study by Lambe which argued that large theropods were pure scavengers because ''Gorgosaurus'' teeth showed hardly any wear. 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. 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. He has put forward arguments in the popular literature to support the pure scavenger hypothesis: * 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. 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 The wolf (''Canis lupus''; : wolves), also known as the grey wolf or gray wolf, is a canine native to Eurasia and North America. More than thirty subspecies of ''Canis lupus'' have been recognized, including the dog and dingo, though gr ...
, hyenas, and secretary birds as well as other extinct animals thought to be predators that would not have used their forelimbs such as phorusrhacids. * Tyrannosaurs had large
olfactory bulb The olfactory bulb (Latin: ''bulbus olfactorius'') is a neural structure of the vertebrate forebrain involved in olfaction, the sense of smell. It sends olfactory information to be further processed in the amygdala, the orbitofrontal cortex (OF ...
s and
olfactory nerve The olfactory nerve, also known as the first cranial nerve, cranial nerve I, or simply CN I, is a cranial nerve that contains sensory nerve fibers relating to the sense of smell. The afferent nerve fibers of the olfactory receptor neurons t ...
s (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 dinosaurs. * Tyrannosaur teeth could crush bone, and therefore could extract as much food (
bone marrow Bone marrow is a semi-solid biological tissue, tissue found within the Spongy bone, spongy (also known as cancellous) portions of bones. In birds and mammals, bone marrow is the primary site of new blood cell production (or haematopoiesis). It i ...
) as possible from carcass remnants, usually the least nutritious parts. Karen Chin and colleagues have found bone fragments in
coprolite A coprolite (also known as a coprolith) is fossilized feces. Coprolites are classified as trace fossils as opposed to body fossils, as they give evidence for the animal's behaviour (in this case, diet) rather than morphology. The name ...
s (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. Summary at * 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. 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. 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 Binocular vision is seeing with two eyes. The Field_of_view, field of view that can be surveyed with two eyes is greater than with one eye. To the extent that the visual fields of the two eyes overlap, #Depth, binocular depth can be perceived. Th ...
slightly better than that of modern hawks. It is not obvious why
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the Heredity, heritable traits characteristic of a population over generation ...
would have favored this long-term trend if tyrannosaurs had been pure scavengers, which would not have needed the advanced
depth perception Depth perception is the ability to perceive distance to objects in the world using the visual system and visual perception. It is a major factor in perceiving the world in three dimensions. Depth sensation is the corresponding term for non-hum ...
that stereoscopic vision provides. 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
vertebra Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spina ...
e. 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. Despite the consensus that the tail bites were caused by ''Tyrannosaurus'', there has been some evidence to show that they might have been created by other factors. For example, a 2014 study suggested that the tail injuries might have been due to ''Edmontosaurus'' individuals stepping on each other, while another study in 2020 backs up the hypothesis that biomechanical stress is the cause for the tail injuries. There is also evidence for an aggressive interaction between a ''
Triceratops ''Triceratops'' ( ; ) is a genus of Chasmosaurinae, chasmosaurine Ceratopsia, ceratopsian dinosaur that lived during the late Maastrichtian age of the Late Cretaceous Period (geology), period, about 68 to 66 million years ago on the island ...
'' 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. 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. 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. Studies on hadrosaur vertebrae from the Hell Creek Formation that were punctured by the teeth of what appears to be a late-stage juvenile ''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. ''Tyrannosaurus'' may have had infectious
saliva Saliva (commonly referred as spit or drool) is an extracellular fluid produced and secreted by salivary glands in the mouth. In humans, saliva is around 99% water, plus electrolytes, mucus, white blood cells, epithelial cells (from which ...
used to kill its prey, as proposed by William Abler in 1992. Abler observed that the (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 The Komodo dragon (''Varanus komodoensis''), also known as the Komodo monitor, is a large reptile of the monitor lizard family Varanidae that is endemic to the Indonesian islands of Komodo (island), Komodo, Rinca, Flores, Gili Dasami, and Gili ...
was thought to have. 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. ''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.


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. 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. A study from Currie, Horner, Erickson and Longrich in 2010 has been put forward as evidence of cannibalism in the genus ''Tyrannosaurus''. They studied some ''Tyrannosaurus'' specimens with tooth marks in the bones, attributable to the same genus. The tooth marks were identified in the
humerus The humerus (; : humeri) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius (bone), radius and ulna, and consists of three sections. The humeral upper extrem ...
, 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.


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 have protected and fed its young. Crocodilians and birds are often suggested by some paleontologists to be modern analogues for dinosaur parenting. 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.


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 researchers found tendon avulsions only among ''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. 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
bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
s. 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. 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''. A more recent study examining the pathologies concluded that the osseous alteration observed most closely 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. One study of ''Tyrannosaurus'' specimens with tooth marks in the bones attributable to the same genus was presented as evidence of cannibalism. Tooth marks in the
humerus The humerus (; : humeri) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius (bone), radius and ulna, and consists of three sections. The humeral upper extrem ...
, foot bones and metatarsals, may indicate opportunistic scavenging, rather than wounds caused by combat with another ''T. rex''. Other tyrannosaurids may also have practiced cannibalism.


Paleoecology

''Tyrannosaurus'' lived during what is referred to as the Lancian faunal stage (
Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
age) at the end of the Late Cretaceous. ''Tyrannosaurus'' ranged from
Canada Canada is a country in North America. Its Provinces and territories of Canada, ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, making it the world's List of coun ...
in the north to at least New Mexico in the south of
Laramidia Laramidia was an island continent that existed during the Late Cretaceous period (99.6–66 Year#SI prefix multipliers, Ma), when the Western Interior Seaway split the continent of North America in two. In the Mesozoic era, Laramidia was an island ...
. During this time ''
Triceratops ''Triceratops'' ( ; ) is a genus of Chasmosaurinae, chasmosaurine Ceratopsia, ceratopsian dinosaur that lived during the late Maastrichtian age of the Late Cretaceous Period (geology), period, about 68 to 66 million years ago on the island ...
'' was the major herbivore in the northern portion of its range, while the titanosaurian
sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their b ...
'' 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 The subtropical zones or subtropics are geographical zone, geographical and Köppen climate classification, climate zones immediately to the Northern Hemisphere, north and Southern Hemisphere, south of the tropics. Geographically part of the Ge ...
, with a warm and humid climate. The flora consisted mostly of angiosperms, but also included trees like dawn redwood ('' Metasequoia''). Archosaurs discovered from this formation include the
ceratopsia Ceratopsia or Ceratopia ( or ; Ancient Greek, Greek: "horned faces") is a group of herbivore, herbivorous, beaked dinosaurs that thrived in what are now North America, Asia and Europe, during the Cretaceous Period (geology), Period, although ance ...
ns ''
Leptoceratops ''Leptoceratops'' (meaning 'small horn face') is a genus of ceratopsian dinosaur from the Late Cretaceous of North America. First found in Alberta in 1910, the type species ''Leptoceratops gracilis'' was named in 1914 by Barnum Brown for a parti ...
'', '' Torosaurus'', and ''
Triceratops ''Triceratops'' ( ; ) is a genus of Chasmosaurinae, chasmosaurine Ceratopsia, ceratopsian dinosaur that lived during the late Maastrichtian age of the Late Cretaceous Period (geology), period, about 68 to 66 million years ago on the island ...
'', the hadrosaurid '' Edmontosaurus annectens'', the parksosaurid '' Thescelosaurus'', the ankylosaurs '' Ankylosaurus'' and '' Denversaurus'', the pachycephalosaurs '' Pachycephalosaurus'' and '' Sphaerotholus'', the paravian theropods '' Acheroraptor'' and '' Pectinodon'', other maniraptoran theropods '' Anzu'', '' Eoneophron'', '' Trierarchuncus'' and indeterminate ornithomimids, and the
pterosaur Pterosaurs are an extinct clade of flying reptiles in the order Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 million to 66 million years ago). Pterosaurs are the earli ...
'' Infernodrakon''. 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 ''Leptoceratops'' (meaning 'small horn face') is a genus of ceratopsian dinosaur from the Late Cretaceous of North America. First found in Alberta in 1910, the type species ''Leptoceratops gracilis'' was named in 1914 by Barnum Brown for a parti ...
'' also lived in the area. In its southern range, specifically based on remains discovered from the North Horn Formation of
Utah Utah is a landlocked state in the Mountain states, Mountain West subregion of the Western United States. It is one of the Four Corners states, sharing a border with Arizona, Colorado, and New Mexico. It also borders Wyoming to the northea ...
, ''Tyrannosaurus rex'' lived alongside the titanosaur '' Alamosaurus'', the ceratopsid '' Torosaurus'' and the indeterminate troodontids and
hadrosaurid Hadrosaurids (), also hadrosaurs or duck-billed dinosaurs, are members of the ornithischian family Hadrosauridae. This group is known as the duck-billed dinosaurs for the flat duck-bill appearance of the bones in their snouts. The ornithopod fami ...
s. ''Tyrannosaurus mcraeensis'' from the McRae Group of
New Mexico New Mexico is a state in the Southwestern United States, Southwestern region of the United States. It is one of the Mountain States of the southern Rocky Mountains, sharing the Four Corners region with Utah, Colorado, and Arizona. It also ...
coexisted with the ceratopsid '' Sierraceratops'' and possibly the titanosaur ''Alamosaurus''. Potential remains identified as cf. ''Tyrannosaurus'' have also been discovered from the Javelina Formation of
Texas Texas ( , ; or ) is the most populous U.S. state, state in the South Central United States, South Central region of the United States. It borders Louisiana to the east, Arkansas to the northeast, Oklahoma to the north, New Mexico to the we ...
, where the remains of the titanosaur ''Alamosaurus'', the ceratopsid '' Bravoceratops'', the
pterosaur Pterosaurs are an extinct clade of flying reptiles in the order Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 million to 66 million years ago). Pterosaurs are the earli ...
s ''
Quetzalcoatlus ''Quetzalcoatlus'' () is a genus of azhdarchid pterosaur that lived during the Maastrichtian age of the Late Cretaceous in North America. The Type (biology), type specimen, recovered in 1971 from the Javelina Formation of Texas, United States, ...
'' and '' Wellnhopterus'', and possible species of troodontids and hadrosaurids are found. Its southern range is thought to have been dominated by semi-arid inland plains, following the probable retreat of the
Western Interior Seaway The Western Interior Seaway (also called the Cretaceous Seaway, the Niobraran Sea, the North American Inland Sea, or the Western Interior Sea) was a large inland sea (geology), inland sea that existed roughly over the present-day Great Plains of ...
as global sea levels fell. ''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. It is possible that tyrannosaurs were originally Asian species, migrating to North America before the end of the Cretaceous period.


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 lower than what should be expected, especially when factoring in that disease pandemics could easily wipe out 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. 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 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. 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. 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. Text was copied from this source, which is available under
Creative Commons Attribution 4.0 International License
.


Cultural significance

Since it was first described in 1905, ''T. rex'' has become the most widely recognized dinosaur species in
popular culture Popular culture (also called pop culture or mass culture) is generally recognized by members of a society as a set of cultural practice, practices, beliefs, artistic output (also known as popular art
f. pop art F is the sixth letter of the Latin alphabet. F may also refer to: Science and technology Mathematics * F or f, the number 15 (number), 15 in hexadecimal and higher positional systems * ''p'F'q'', the hypergeometric function * F-distributi ...
or mass art, sometimes contraste ...
. 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. Robert T. Bakker notes this in '' The Dinosaur Heresies'' and explains that, "a name like ''T. rex'' is just irresistible to the tongue."


See also

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


Notes


References


Further reading

*


External links


The University of Edinburgh Lecture Dr Stephen Brusatte – Tyrannosaur Discoveries Feb 20, 2015


Stephen Brusatte Thomas Carr 2016
Australia's answer to T-Rex
State Library of Queensland State Library of Queensland (State Library) is the state public reference and research library of Queensland, Australia, operated by the Government of Queensland, state government. The Library is governed by the Library Board of Queensland, whi ...


Exhibits


American Museum of Natural History
{{Authority control Dinosaur genera Maastrichtian dinosaurs Hell Creek Formation Lance Formation Laramie Formation Scollard Formation Taxa named by Henry Fairfield Osborn Fossil taxa described in 1905 Dinosaurs of Canada Dinosaurs of the United States Apex predators