Dinosaur

Dinosaurs are a diverse group of reptiles[note 1] of the clade Dinosauria.

They first appeared during the Triassic period, between 245 and Dinosaurs
233.23  million years ago (mya), although the exact origin and timing of the Temporal range: (Possible Middle
evolution of dinosaurs is a subject of active research. They became the Triassic record)
dominant terrestrial vertebrates after the Triassic–Jurassic extinction event
201.3  mya and their dominance continued throughout the Jurassic and
Cretaceous periods. The fossil record shows that birds are feathered dinosaurs,
having evolved from earlier theropods during the Late Jurassic epoch, and are
the only dinosaur lineage known to have survived the Cretaceous–Paleogene
extinction event approximately 66  mya. Dinosaurs can therefore be divided
into avian dinosaurs—birds—and the extinct non-avian dinosaurs, which Stegosaurus stenops
are all dinosaurs other than birds. (a stegosaur)

Dinosaurs are varied from taxonomic, morphological and ecological

standpoints. Birds, at over 10,700 living species, are among the most diverse
groups of vertebrates. Using fossil evidence, paleontologists have identified
over 900 distinct genera and more than 1,000 different species of non-avian
dinosaurs. Dinosaurs are represented on every continent by both extant species
Herrerasaurus
(birds) and fossil remains. Through the first half of the 20th century, before
ischigualastensis
birds were recognized as dinosaurs, most of the scientific community believed
(a carnivorous basal
dinosaurs to have been sluggish and cold-blooded. Most research conducted
dinosaur)
since the 1970s, however, has indicated that dinosaurs were active animals
with elevated metabolisms and numerous adaptations for social interaction.
Some were herbivorous, others carnivorous. Evidence suggests that all
dinosaurs were egg-laying, and that nest-building was a trait shared by many
dinosaurs, both avian and non-avian.

While dinosaurs were ancestrally bipedal, many extinct groups included Triceratops horridus
quadrupedal species, and some were able to shift between these stances. (a ceratopsian)
Elaborate display structures such as horns or crests are common to all dinosaur
groups, and some extinct groups developed skeletal modifications such as
bony armor and spines. While the dinosaurs' modern-day surviving avian
lineage (birds) are generally small due to the constraints of flight, many
prehistoric dinosaurs (non-avian and avian) were large-bodied—the largest
sauropod dinosaurs are estimated to have reached lengths of 39.7 meters (130
Apatosaurus louisae
feet) and heights of 18 m (59 ft) and were the largest land animals of all time.
(a sauropod)
The misconception that non-avian dinosaurs were uniformly gigantic is based
in part on preservation bias, as large, sturdy bones are more likely to last until
they are fossilized. Many dinosaurs were quite small, some measuring about
50 centimeters (20 inches) in length.
Edmontosaurus regalis
The first dinosaur fossils were recognized in the early 19th century, with the (a hadrosaurid
name "dinosaur" (meaning "terrible lizard") being coined by Sir Richard ornithopod)
Owen in 1842 to refer to these "great fossil lizards".[7][8][9] Since then,
mounted fossil dinosaur skeletons have been major attractions at museums
worldwide, and dinosaurs have become an enduring part of popular culture.
The large sizes of some dinosaurs, as well as their seemingly monstrous and
fantastic nature, have ensured their regular appearance in best-selling books Microraptor gui
(a dromaeosaurid
theropod)
and films, such as Jurassic Park. Persistent public enthusiasm for the animals Scientific classification
has resulted in significant funding for dinosaur science, and new discoveries
are regularly covered by the media. Domain: Eukaryota
Kingdom: Animalia
Definition Phylum: Chordata

Under phylogenetic nomenclature, dinosaurs are usually defined as the group Clade: Sauropsida
consisting of the most recent common ancestor (MRCA) of Triceratops and Clade: Archosauria
modern birds (Neornithes), and all its descendants.[10] It has also been
suggested that Dinosauria be defined with respect to the MRCA of Clade: Avemetatarsalia
Megalosaurus and Iguanodon, because these were two of the three genera Clade: Ornithodira
cited by Richard Owen when he recognized the Dinosauria.[11] Both
Clade: Dinosauromorpha
definitions result in the same set of animals being defined as dinosaurs:
"Dinosauria = Ornithischia + Saurischia". This definition includes major Clade: Dinosauriformes
groups such as ankylosaurians (armored herbivorous quadrupeds),
Clade: Dracohors
stegosaurians (plated herbivorous quadrupeds), ceratopsians (bipedal or
quadrupedal herbivores with neck frills), pachycephalosaurians (bipedal Clade: Dinosauria
herbivores with thick skulls), ornithopods (bipedal or quadrupedal herbivores Owen, 1842
including "duck-bills"), theropods (mostly bipedal carnivores and birds), and
sauropodomorphs (mostly large herbivorous quadrupeds with long necks and Major groups
tails).[12]
†Ornithischia
Birds are the sole surviving dinosaurs. In traditional taxonomy, birds were
considered a separate class that had evolved from dinosaurs, a distinct †Sauropodomorpha
superorder. However, a majority of contemporary paleontologists concerned Theropoda
with dinosaurs reject the traditional style of classification in favor of
phylogenetic taxonomy; this approach requires that, for a group to be natural, Various extinct groups
all descendants of members of the group must be included in the group.[13] Aves (birds)
Birds belong to the dinosaur subgroup Maniraptora, which are coelurosaurs,
which are theropods, which are saurischians.[14] Possible dinosaurs of

Research by Matthew G. Baron, David B. Norman, and Paul M. Barrett in uncertain affinity
2017 suggested a radical revision of dinosaurian systematics. Phylogenetic
analysis by Baron et al. recovered the Ornithischia as being closer to the †Alwalkeria?
Theropoda than the Sauropodomorpha, as opposed to the traditional union of †Chilesaurus
theropods with sauropodomorphs. This would cause sauropods and kin to fall
outside traditional dinosaurs, so they re-defined Dinosauria as the last common †Chindesaurus?
ancestor of Triceratops horridus, Passer domesticus and Diplodocus carnegii, †Daemonosaurus?
and all of its descendants, to ensure that sauropods and kin remain included as
†Eodromaeus?
dinosaurs. They also resurrected the clade Ornithoscelida to refer to the group
containing Ornithischia and Theropoda.[15][16] †Nhandumirim
†Nyasasaurus?
General description †Pisanosaurus?
†Smok?
Using one of the above definitions, dinosaurs can be generally described as
archosaurs with hind limbs held erect beneath the body.[17] Other prehistoric †Tawa?
animals, including pterosaurs, mosasaurs, ichthyosaurs, plesiosaurs, and †Thecospondylus
Dimetrodon, while often popularly conceived of as dinosaurs, are not
taxonomically classified as dinosaurs.[18] Pterosaurs are distantly related to †Guaibasauridae?
dinosaurs, being members of the clade Ornithodira. The other groups †Herrerasauria?[1][2]
mentioned are, like dinosaurs and pterosaurs, members of Sauropsida (the
reptile and bird clade), except Dimetrodon (which is a synapsid). None of †Silesauridae?
them had the erect hind limb posture characteristic of true dinosaurs.[19] (paraphyletic?) [3][4][5][6]

Dinosaurs were the dominant terrestrial vertebrates of the Mesozoic Era,

especially the Jurassic and Cretaceous periods. Other groups of animals were restricted in size and niches; mammals, for
example, rarely exceeded the size of a domestic cat, and were generally rodent-sized carnivores of small prey.[20] They
 have always been recognized as an
extremely varied group of animals;
over 900 non-avian dinosaur genera
have been identified with certainty as
of 2018, and the total number of
genera preserved in the fossil record
has been estimated at around 1850,
nearly 75% of which remain to be
discovered, and 1124 species by
2016.[21][22][23] A 1995 study
predicted that about 3,400 dinosaur
Triceratops skeleton, Natural History
genera ever existed, including many
Museum of Los Angeles County Birds are avian dinosaurs, and in
that would not have been preserved
phylogenetic taxonomy are included
in the fossil record.[24]
in the group Dinosauria.
In 2016, the estimated number of dinosaur species that existed in the Mesozoic
was 1,543–2,468.[25][26] In 2021, the number of modern-day birds (avian
dinosaurs) was estimated to be at 10,806 species.[27] Some are herbivorous, others carnivorous, including seed-eaters,
fish-eaters, insectivores, and omnivores. While dinosaurs were ancestrally bipedal (as are all modern birds), some
prehistoric species were quadrupeds, and others, such as Anchisaurus and Iguanodon, could walk just as easily on two or
four legs. Cranial modifications like horns and crests are common dinosaurian traits, and some extinct species had bony
armor. Although known for large size, many Mesozoic dinosaurs were human-sized or smaller, and modern birds are
generally small in size. Dinosaurs today inhabit every continent, and fossils show that they had achieved global
distribution by at least the Early Jurassic epoch.[28] Modern birds inhabit most available habitats, from terrestrial to
marine, and there is evidence that some non-avian dinosaurs (such as Microraptor) could fly or at least glide, and others,
such as spinosaurids, had semiaquatic habits.[29]

Distinguishing anatomical features

While recent discoveries have made it more difficult to present a universally agreed-upon list of their distinguishing
features, nearly all dinosaurs discovered so far share certain modifications to the ancestral archosaurian skeleton, or are
clearly descendants of older dinosaurs showing these modifications. Although some later groups of dinosaurs featured
further modified versions of these traits, they are considered typical for Dinosauria; the earliest dinosaurs had them and
passed them on to their descendants. Such modifications, originating in the most recent common ancestor of a certain
taxonomic group, are called the synapomorphies of such a group.[30]

A detailed assessment of archosaur interrelations by Sterling Nesbitt[31]

confirmed or found the following twelve unambiguous synapomorphies, some
previously known:

In the skull, a supratemporal fossa (excavation) is present in front of

the supratemporal fenestra, the main opening in the rear skull roof
Epipophyses, obliquely backward-pointing processes on the rear top
corners of the anterior (front) neck vertebrae behind the atlas and Labeled diagram of a typical
axis, the first two neck vertebrae archosaur skull, the skull of
Apex of a deltopectoral crest (a projection on which the deltopectoral Dromaeosaurus
muscles attach) located at or more than 30% down the length of the
humerus (upper arm bone)
Radius, a lower arm bone, shorter than 80% of humerus length
Fourth trochanter (projection where the caudofemoralis muscle attaches on the inner rear shaft) on the
femur (thigh bone) is a sharp flange
Fourth trochanter asymmetrical, with distal, lower, margin forming a steeper angle to the shaft
On the astragalus and calcaneum, upper ankle bones, the proximal articular facet, the top connecting
surface, for the fibula occupies less than 30% of the transverse width of the element
Exoccipitals (bones at the back of the skull) do not meet along the midline on the floor of the endocranial
cavity, the inner space of the braincase
 In the pelvis, the proximal articular surfaces of the ischium with the ilium and the pubis are separated by
a large concave surface (on the upper side of the ischium a part of the open hip joint is located between
the contacts with the pubic bone and the ilium)
Cnemial crest on the tibia (protruding part of the top surface of the shinbone) arcs anterolaterally (curves
to the front and the outer side)
Distinct proximodistally oriented (vertical) ridge present on the posterior face of the distal end of the tibia
(the rear surface of the lower end of the shinbone)
Concave articular surface for the fibula of the calcaneum (the top surface of the calcaneum, where it
touches the fibula, has a hollow profile)

Nesbitt found a number of further potential synapomorphies and discounted a number of synapomorphies previously
suggested. Some of these are also present in silesaurids, which Nesbitt recovered as a sister group to Dinosauria,
including a large anterior trochanter, metatarsals II and IV of subequal length, reduced contact between ischium and
pubis, the presence of a cnemial crest on the tibia and of an ascending process on the astragalus, and many others.[10]

A variety of other skeletal features are shared by dinosaurs. However, because

they either are common to other groups of archosaurs or were not present in all
early dinosaurs, these features are not considered to be synapomorphies. For
example, as diapsids, dinosaurs ancestrally had two pairs of Infratemporal
fenestrae (openings in the skull behind the eyes), and as members of the diapsid
group Archosauria, had additional openings in the snout and lower jaw.[32]
Hip joints and hindlimb postures of:
Additionally, several characteristics once thought to be synapomorphies are now
(left to right) typical reptiles
known to have appeared before dinosaurs, or were absent in the earliest dinosaurs
(sprawling), dinosaurs and mammals
and independently evolved by different dinosaur groups. These include an
(erect), and rauisuchians (pillar-
elongated scapula, or shoulder blade; a sacrum composed of three or more fused
erect)
vertebrae (three are found in some other archosaurs, but only two are found in
Herrerasaurus);[10] and a perforate acetabulum, or hip socket, with a hole at the
center of its inside surface (closed in Saturnalia tupiniquim, for example).[33][34] Another difficulty of determining
distinctly dinosaurian features is that early dinosaurs and other archosaurs from the Late Triassic epoch are often poorly
known and were similar in many ways; these animals have sometimes been misidentified in the literature.[35]

Dinosaurs stand with their hind limbs erect in a manner similar to most modern mammals, but distinct from most other
reptiles, whose limbs sprawl out to either side.[36] This posture is due to the development of a laterally facing recess in
the pelvis (usually an open socket) and a corresponding inwardly facing distinct head on the femur.[37] Their erect
posture enabled early dinosaurs to breathe easily while moving, which likely permitted stamina and activity levels that
surpassed those of "sprawling" reptiles.[38] Erect limbs probably also helped support the evolution of large size by
reducing bending stresses on limbs.[39] Some non-dinosaurian archosaurs, including rauisuchians, also had erect limbs
but achieved this by a "pillar-erect" configuration of the hip joint, where instead of having a projection from the femur
insert on a socket on the hip, the upper pelvic bone was rotated to form an overhanging shelf.[39]

History of study

Pre-scientific history

Dinosaur fossils have been known for millennia, although their true nature was not recognized. The Chinese considered
them to be dragon bones and documented them as such. For example, Huayang Guo Zhi  ( 華陽國志 ), a gazetteer
compiled by Chang Qu ( 常璩 ) during the Western Jin Dynasty (265–316), reported the discovery of dragon bones at
Wucheng in Sichuan Province.[40] Villagers in central China have long unearthed fossilized "dragon bones" for use in
traditional medicines.[41] In Europe, dinosaur fossils were generally believed to be the remains of giants and other
biblical creatures.[42]

Early dinosaur research

 Scholarly descriptions of what would now be recognized as dinosaur bones first appeared
in the late 17th century in England. Part of a bone, now known to have been the femur of
a Megalosaurus,[43] was recovered from a limestone quarry at Cornwell near Chipping
Norton, Oxfordshire, in 1676. The fragment was sent to Robert Plot, Professor of
Chemistry at the University of Oxford and first curator of the Ashmolean Museum, who
published a description in his The Natural History of Oxford-shire (1677).[44] He
correctly identified the bone as the lower extremity of the femur of a large animal, and
recognized that it was too large to belong to any known species. He, therefore, concluded
it to be the femur of a huge human, perhaps a Titan or another type of giant featured in
legends.[45][46] Edward Lhuyd, a friend of Sir Isaac Newton, published Lithophylacii
Britannici ichnographia (1699), the first scientific treatment of what would now be
recognized as a dinosaur when he described and named a sauropod tooth, "Rutellum
William Buckland
impicatum",[47][48] that had been found in Caswell, near Witney, Oxfordshire.[49]

Between 1815 and 1824, the Rev William

Buckland, the first Reader of Geology at the University of Oxford, collected
more fossilized bones of Megalosaurus and became the first person to describe a
non-avian dinosaur in a scientific journal.[43][50] The second non-avian dinosaur
genus to be identified, Iguanodon, was discovered in 1822 by Mary Ann
Mantell  – the wife of English geologist Gideon Mantell. Gideon Mantell
recognized similarities between his fossils and the bones of modern iguanas. He
published his findings in 1825.[51][52]

The study of these "great fossil lizards" soon became of great interest to
European and American scientists, and in 1842 the English paleontologist Sir
Richard Owen coined the term "dinosaur", using it to refer to the "distinct tribe or Sir Richard Owen's coining of the
sub-order of Saurian Reptiles" that were then being recognized in England and word dinosaur, in the 1842 revised
around the world.[7][8][9][53][54] The term is derived from Ancient Greek δεινός version of his talk at an 1841
(deinos)  'terrible, potent or fearfully great', and σαῦρος (sauros)  'lizard or meeting of the British Association for
reptile'.[53][55] Though the taxonomic name has often been interpreted as a the Advancement of Science.
reference to dinosaurs' teeth, claws, and other fearsome characteristics, Owen
intended it also to evoke their size and majesty.[56] Owen recognized that the
remains that had been found so far, Iguanodon, Megalosaurus and Hylaeosaurus, shared distinctive features, and so
decided to present them as a distinct taxonomic group. As clarified by British geologist and historian Hugh Torrens,
Owen had given a presentation about fossil reptiles to the British Association for the Advancement of Science in 1841,
but reports of the time show that Owen did not mention the word "dinosaur", nor recognize dinosaurs as a distinct group
of reptiles in his address. He introduced the Dinosauria only in the revised text version of his talk published in April of
1842.[7][8] With the backing of Prince Albert, the husband of Queen Victoria, Owen established the Natural History
Museum, London, to display the national collection of dinosaur fossils and other biological and geological exhibits.[57]

Discoveries in North America

In 1858, William Parker Foulke discovered the first known American

dinosaur, in marl pits in the small town of Haddonfield, New Jersey.
(Although fossils had been found before, their nature had not been
correctly discerned.) The creature was named Hadrosaurus foulkii. It
was an extremely important find: Hadrosaurus was one of the first
nearly complete dinosaur skeletons found (the first was in 1834, in
Maidstone, England), and it was clearly a bipedal creature. This was a
revolutionary discovery as, until that point, most scientists had
believed dinosaurs walked on four feet, like other lizards. Foulke's
Edward Drinker Cope Othniel Charles Marsh discoveries sparked a wave of interests in dinosaurs in the United
States, known as dinosaur mania.[58]

Dinosaur mania was exemplified by the fierce rivalry between Edward Drinker Cope and Othniel Charles Marsh, both of
whom raced to be the first to find new dinosaurs in what came to be known as the Bone Wars. This fight between the
two scientists lasted for over 30 years, ending in 1897 when Cope died after spending his entire fortune on the dinosaur
hunt. Many valuable dinosaur specimens were damaged or destroyed due to the pair's rough methods: for example, their
diggers often used dynamite to unearth bones. Modern paleontologists would find such methods crude and unacceptable,
since blasting easily destroys fossil and stratigraphic evidence. Despite their unrefined methods, the contributions of Cope
and Marsh to paleontology were vast: Marsh unearthed 86 new species of dinosaur and Cope discovered 56, a total of
142 new species. Cope's collection is now at the American Museum of Natural History in New York City, while Marsh's
is at the Peabody Museum of Natural History at Yale University.[59]

"Dinosaur renaissance" and beyond

World War II caused a pause in palaeontological research; after the war, research
attention was also diverted increasingly to fossil mammals rather than dinosaurs,
which were seen as sluggish and cold-blooded.[60][61] At the end of the 1960s,
however, the field of dinosaur research experienced a surge in activity that
remains ongoing.[62] Several seminal studies led to this activity. First, John John Ostrom's original restoration of
Ostrom discovered the bird-like dromaeosaurid theropod Deinonychus and Deinonychus, published in 1969
described it in 1969. Its anatomy indicated that it was an active predator that was
likely warm-blooded, in marked contrast to the then-prevailing image of
dinosaurs.[60] Concurrently, Robert T. Bakker published a series of studies that likewise argued for active lifestyles in
dinosaurs based on anatomical and ecological evidence (see § Physiology),[63][64] which were subsequently summarized
in his 1986 book The Dinosaur Heresies.[65]

New revelations were supported by an increase in dinosaur discoveries. Major

new dinosaur discoveries have been made by paleontologists working in
previously unexplored regions, including India, South America, Madagascar,
Antarctica, and most significantly China. Across theropods, sauropodomorphs,
and ornithischians, the number of named genera began to increase exponentially
in the 1990s.[21] As of 2008, over 30 new species of dinosaurs were named each
year.[66] At least sauropodomorphs experienced a further increase in the number
of named species in the 2010s, with an average of 9.3 new species having been
Paleontologist Robert T. Bakker with named each year between 2009 and 2020. As a consequence, more
a mounted skeleton of a sauropodomorphs were named between 1990 and 2020 than in all previous years
tyrannosaurid (Gorgosaurus libratus) combined.[67] These new localities also led to improvements in overall specimen
quality, with new species being increasingly named not on scrappy fossils but on
more complete skeletons, sometimes from multiple individuals. Better specimens
also led to new species being invalidated less frequently.[66] Asian localities have produced the most complete theropod
specimens,[68] while North American localities have produced the most complete sauropodomorph specimens.[67]

Prior to the dinosaur renaissance, dinosaurs were mostly classified using the traditional rank-based system of Linnaean
taxonomy. The renaissance was also accompanied by the increasingly widespread application of cladistics, a more
objective method of classification based on ancestry and shared traits, which has proved tremendously useful in the study
of dinosaur systematics and evolution. Cladistic analysis, among other techniques, helps to compensate for an often
incomplete and fragmentary fossil record.[69][70] Reference books summarizing the state of dinosaur research, such as
David B. Weishampel and colleagues' The Dinosauria, made knowledge more accessible[71] and spurred further interest
in dinosaur research. The release of the first and second editions of The Dinosauria in 1990 and 2004, and of a review
paper by Paul Sereno in 1998, were accompanied by increases in the number of published phylogenetic trees for
dinosaurs.[72]

Soft tissue and molecular preservation

Dinosaur fossils are not limited to bones, but also include imprints or mineralized remains of skin coverings, organs, and
other tissues. Of these, skin coverings based on keratin proteins are most easily preserved because of their cross-linked,
hydrophobic molecular structure.[73] Fossils of keratin-based skin coverings or bony skin coverings are known from
most major groups of dinosaurs. Dinosaur fossils with scaly skin impressions have been found since the 19th century.
Samuel Beckles discovered a sauropod forelimb with preserved skin in 1852 that was incorrectly attributed to a
crocodile; it was correctly attributed by Marsh in 1888 and subject to further study by Reginald Hooley in 1917.[74]
Among ornithischians, in 1884 Jacob Wortman found skin impressions on the first known specimen of Edmontosaurus
annectens, which were largely destroyed during the specimen's excavation.[75]
Owen and Hooley subsequently described skin impressions of Hypsilophodon
and Iguanodon in 1885 and 1917.[74] Since then, scale impressions have been
most frequently found among hadrosaurids, where the impressions are known
from nearly the entire body across multiple specimens.[76]

Starting from the 1990s, major discoveries

of exceptionally preserved fossils in deposits
known as conservation Lagerstätten An Edmontosaurus specimen's skin
contributed to research on dinosaur soft impressions found in 1999
Colour restoration of
tissues.[77][78] Chiefly among these were the
Sinosauropteryx
rocks that produced the Jehol (Early
Cretaceous) and Yanliao (Mid-to-Late Jurassic) biotas of northeastern China, from
which hundreds of dinosaur specimens bearing impressions of feather-like structures
(both closely related to birds and otherwise, see §  Origin of birds) have been
described by Xing Xu and colleagues.[79][80] In living reptiles and mammals,
pigment-storing cellular structures known as melanosomes are partially responsible
Colour restoration of
Psittacosaurus
for producing colouration.[81][82] Both chemical traces of melanin and
characteristically-shaped melanosomes have been reported from feathers and scales
of Jehol and Yanliao dinosaurs, including both theropods and ornithischians.[83]
This has enabled multiple full-body reconstructions of dinosaur colouration, such as for Sinosauropteryx[84] and
Psittacosaurus[85] by Jakob Vinther and colleagues, and similar techniques have also been extended to dinosaur fossils
from other localities.[81] (However, some researchers have also suggested that fossilized melanosomes represent bacterial
remains.[86][87]) Stomach contents in some Jehol and Yanliao dinosaurs closely related to birds have also provided
indirect indications of diet and digestive system anatomy (e.g., crops).[88][89] More concrete evidence of internal anatomy
has been reported in Scipionyx from the Pietraroja Plattenkalk of Italy. It preserves portions of the intestines, colon, liver,
muscles, and windpipe.[90]

Concurrently, a line of work led by Mary Higby Schweitzer, Jack Horner, and
colleagues reported various occurrences of preserved soft tissues and proteins
within dinosaur bone fossils. Various mineralized structures that likely
represented red blood cells and collagen fibres had been found by Schweitzer and
others in tyrannosaurid bones as early as 1991.[91][92][93] However, in 2005,
Schweitzer and colleagues reported that a femur of Tyrannosaurus preserved soft,
flexible tissue within, including blood vessels, bone matrix, and connective tissue
(bone fibers) that had retained their microscopic structure.[94] This discovery
suggested that original soft tissues could be preserved over geological time,[73] Scipionyx fossil with intestines,
with multiple mechanisms having been proposed.[95] Later, in 2009, Schweitzer Natural History Museum of Milan
and colleagues reported that a Brachylophosaurus femur preserved similar
microstructures, and immunohistochemical techniques (based on antibody
binding) demonstrated the presence of proteins such as collagen, elastin, and laminin.[96] Both specimens yielded
collagen protein sequences that were viable for molecular phylogenetic analyses, which grouped them with birds as
would be expected.[96][97] The extraction of fragmentary DNA has also been reported for both of these fossils,[98] along
with a specimen of Hypacrosaurus.[99] In 2015, Sergio Bertazzo and colleagues reported the preservation of collagen
fibres and red blood cells in eight Cretaceous dinosaur specimens that did not show any signs of exceptional
preservation, indicating that soft tissue may be preserved more commonly than previously thought.[100] Suggestions that
these structures represent bacterial biofilms[101] have been rejected,[102] but cross-contamination remains a possibility
that is difficult to detect.[103]

Evolutionary history

Origins and early evolution

Dinosaurs diverged from their archosaur ancestors during the Middle to Late
Triassic epochs, roughly 20  million years after the devastating Permian–Triassic
extinction event wiped out an estimated 96% of all marine species and 70% of
terrestrial vertebrate species approximately 252  million years ago.[104][105] The
oldest dinosaur fossils known from substantial remains date to the Carnian epoch
of the Triassic period and have been found primarily in the Ischigualasto and
Santa Maria Formations of Argentina, and the Pebbly Arkose Formation of
Zimbabwe.[106]
The early dinosaurs Herrerasaurus
The Ischigualasto Formation (radiometrically dated at 231-230 million years (large), Eoraptor (small) and a
old[107]) has produced the early saurischian Eoraptor, originally considered a Plateosaurus skull, from the Triassic
member of the Herrerasauridae[108] but now considered to be an early
sauropodomorph, along with the herrerasaurids Herrerasaurus and
Sanjuansaurus, and the sauropodomorphs Chromogisaurus, Eodromaeus, and Panphagia.[109] Eoraptor's likely
resemblance to the common ancestor of all dinosaurs suggests that the first dinosaurs would have been small, bipedal
predators.[110][111][112] The Santa Maria Formation (radiometrically dated to be older, at 233.23 million years old[113])
has produced the herrerasaurids Gnathovorax and Staurikosaurus, along with the sauropodomorphs Bagualosaurus,
Buriolestes, Guaibasaurus, Macrocollum, Nhandumirim, Pampadromaeus, Saturnalia, and Unaysaurus.[109] The
Pebbly Arkose Formation, which is of uncertain age but was likely comparable to the other two, has produced the
sauropodomorph Mbiresaurus, along with an unnamed herrerasaurid.[106]

Less well-preserved remains of the sauropodomorphs Jaklapallisaurus and Nambalia, along with the early saurischian
Alwalkeria, are known from the Upper Maleri and Lower Maleri Formations of India.[114] The Carnian-aged Chañares
Formation of Argentina preserves primitive, dinosaur-like ornithodirans such as Lagosuchus and Lagerpeton in
Argentina, making it another important site for understanding dinosaur evolution. These ornithodirans support the model
of early dinosaurs as small, bipedal predators.[109][115] Dinosaurs may have appeared as early as the Anisian epoch of
the Triassic, approximately 245  million years ago, which is the age of Nyasasaurus from the Manda Formation of
Tanzania. However, its known fossils are too fragmentary to identify it as a dinosaur or only a close relative.[116] The
referral of the Manda Formation to the Anisian is also uncertain. Regardless, dinosaurs existed alongside non-dinosaurian
ornithodirans for a period of time, with estimates ranging from 5–10 million years[117] to 21 million years.[113]

When dinosaurs appeared, they were not the dominant terrestrial animals. The terrestrial habitats were occupied by
various types of archosauromorphs and therapsids, like cynodonts and rhynchosaurs. Their main competitors were the
pseudosuchians, such as aetosaurs, ornithosuchids and rauisuchians, which were more successful than the dinosaurs.[118]
Most of these other animals became extinct in the Triassic, in one of two events. First, at about 215 million years ago, a
variety of basal archosauromorphs, including the protorosaurs, became extinct. This was followed by the Triassic–
Jurassic extinction event (about 201 million years ago), that saw the end of most of the other groups of early archosaurs,
like aetosaurs, ornithosuchids, phytosaurs, and rauisuchians. Rhynchosaurs and dicynodonts survived (at least in some
areas) at least as late as early –mid Norian and late Norian or earliest Rhaetian stages, respectively,[119][120] and the exact
date of their extinction is uncertain. These losses left behind a land fauna of crocodylomorphs, dinosaurs, mammals,
pterosaurians, and turtles.[10] The first few lines of early dinosaurs diversified through the Carnian and Norian stages of
the Triassic, possibly by occupying the niches of the groups that became extinct.[12] Also notably, there was a heightened
rate of extinction during the Carnian pluvial event.[121]

Evolution and paleobiogeography

Dinosaur evolution after the Triassic followed changes in

vegetation and the location of continents. In the Late Triassic
and Early Jurassic, the continents were connected as the single
landmass Pangaea, and there was a worldwide dinosaur fauna
mostly composed of coelophysoid carnivores and early
sauropodomorph herbivores.[122] Gymnosperm plants
(particularly conifers), a potential food source, radiated in the
Late Triassic. Early sauropodomorphs did not have
sophisticated mechanisms for processing food in the mouth, and The supercontinent Pangaea in the early Mesozoic
so must have employed other means of breaking down food (around 200 million years ago)
farther along the digestive tract.[123] The general homogeneity
of dinosaurian faunas continued into the Middle and Late Jurassic, where most localities had predators consisting of
ceratosaurians, megalosauroids, and allosauroids, and herbivores consisting of stegosaurian ornithischians and large
sauropods. Examples of this include the Morrison Formation of North America and Tendaguru Beds of Tanzania.
Dinosaurs in China show some differences, with specialized metriacanthosaurid theropods and unusual, long-necked
sauropods like Mamenchisaurus.[122] Ankylosaurians and ornithopods were also becoming more common, but primitive
sauropodomorphs had become extinct. Conifers and pteridophytes were the most common plants. Sauropods, like earlier
sauropodomorphs, were not oral processors, but ornithischians were evolving various means of dealing with food in the
mouth, including potential cheek-like organs to keep food in the mouth, and jaw motions to grind food.[123] Another
notable evolutionary event of the Jurassic was the appearance of true birds, descended from maniraptoran
coelurosaurians.[14]

By the Early Cretaceous and the ongoing breakup of Pangaea, dinosaurs were becoming strongly differentiated by
landmass. The earliest part of this time saw the spread of ankylosaurians, iguanodontians, and brachiosaurids through
Europe, North America, and northern Africa. These were later supplemented or replaced in Africa by large spinosaurid
and carcharodontosaurid theropods, and rebbachisaurid and titanosaurian sauropods, also found in South America. In
Asia, maniraptoran coelurosaurians like dromaeosaurids, troodontids, and oviraptorosaurians became the common
theropods, and ankylosaurids and early ceratopsians like Psittacosaurus became important herbivores. Meanwhile,
Australia was home to a fauna of basal ankylosaurians, hypsilophodonts, and iguanodontians.[122] The stegosaurians
appear to have gone extinct at some point in the late Early Cretaceous or early Late Cretaceous. A major change in the
Early Cretaceous, which would be amplified in the Late Cretaceous, was the evolution of flowering plants. At the same
time, several groups of dinosaurian herbivores evolved more sophisticated ways to orally process food. Ceratopsians
developed a method of slicing with teeth stacked on each other in batteries, and iguanodontians refined a method of
grinding with dental batteries, taken to its extreme in hadrosaurids.[123] Some sauropods also evolved tooth batteries, best
exemplified by the rebbachisaurid Nigersaurus.[124]

There were three general dinosaur faunas in the Late Cretaceous. In the northern continents of North America and Asia,
the major theropods were tyrannosaurids and various types of smaller maniraptoran theropods, with a predominantly
ornithischian herbivore assemblage of hadrosaurids, ceratopsians, ankylosaurids, and pachycephalosaurians. In the
southern continents that had made up the now-splitting supercontinent Gondwana, abelisaurids were the common
theropods, and titanosaurian sauropods the common herbivores. Finally, in Europe, dromaeosaurids, rhabdodontid
iguanodontians, nodosaurid ankylosaurians, and titanosaurian sauropods were prevalent.[122] Flowering plants were
greatly radiating,[123] with the first grasses appearing by the end of the Cretaceous.[125] Grinding hadrosaurids and
shearing ceratopsians became very diverse across North America and Asia. Theropods were also radiating as herbivores
or omnivores, with therizinosaurians and ornithomimosaurians becoming common.[123]

The Cretaceous–Paleogene extinction event, which occurred approximately 66  million years ago at the end of the
Cretaceous, caused the extinction of all dinosaur groups except for the neornithine birds. Some other diapsid groups,
including crocodilians, dyrosaurs, sebecosuchians, turtles, lizards, snakes, sphenodontians, and choristoderans, also
survived the event.[126]

The surviving lineages of neornithine birds, including the ancestors of modern ratites, ducks and chickens, and a variety
of waterbirds, diversified rapidly at the beginning of the Paleogene period, entering ecological niches left vacant by the
extinction of Mesozoic dinosaur groups such as the arboreal enantiornithines, aquatic hesperornithines, and even the
larger terrestrial theropods (in the form of Gastornis, eogruiids, bathornithids, ratites, geranoidids, mihirungs, and "terror
birds"). It is often stated that mammals out-competed the neornithines for dominance of most terrestrial niches but many
of these groups co-existed with rich mammalian faunas for most of the Cenozoic Era.[127] Terror birds and bathornithids
occupied carnivorous guilds alongside predatory mammals,[128][129] and ratites are still fairly successful as mid-sized
herbivores; eogruiids similarly lasted from the Eocene to Pliocene, becoming extinct only very recently after over
20 million years of co-existence with many mammal groups.[130]

Classification
Dinosaurs belong to a group known as archosaurs, which also includes modern crocodilians. Within the archosaur group,
dinosaurs are differentiated most noticeably by their gait. Dinosaur legs extend directly beneath the body, whereas the
legs of lizards and crocodilians sprawl out to either side.[30]
Collectively, dinosaurs as a clade are divided into two primary
branches, Saurischia and Ornithischia. Saurischia includes
those taxa sharing a more recent common ancestor with birds
than with Ornithischia, while Ornithischia includes all taxa
sharing a more recent common ancestor with Triceratops than
with Saurischia. Anatomically, these two groups can be
distinguished most noticeably by their pelvic structure. Early
saurischians—"lizard-hipped", from the Greek sauros
(σαῦρος) meaning "lizard" and ischion (ἰσχίον) meaning "hip
joint"—retained the hip structure of their ancestors, with a
Saurischian pelvis structure Tyrannosaurus pelvis
pubis bone directed cranially, or forward.[37] This basic form
(left side) (showing saurischian
was modified by rotating the pubis backward to varying
structure – left side)
degrees in several groups (Herrerasaurus,[131]
therizinosauroids, [132] dromaeosaurids, [133] and birds[14]).
Saurischia includes the theropods (exclusively bipedal and
with a wide variety of diets) and sauropodomorphs (long-
necked herbivores which include advanced, quadrupedal
groups).[29][134]

By contrast, ornithischians—"bird-hipped", from the Greek

ornitheios (ὀρνίθειος) meaning "of a bird" and ischion Ornithischian pelvis Edmontosaurus pelvis (showing
(ἰσχίον) meaning "hip joint"—had a pelvis that superficially structure (left side) ornithischian structure – left
side)
resembled a bird's pelvis: the pubic bone was oriented
caudally (rear-pointing). Unlike birds, the ornithischian pubis
also usually had an additional forward-pointing process.
Ornithischia includes a variety of species that were primarily herbivores.

Despite the terms "bird hip" (Ornithischia) and "lizard hip" (Saurischia), birds are not part of Ornithischia. Birds instead
belong to Saurischia, the “lizard-hipped” dinosaurs—birds evolved from earlier dinosaurs with "lizard hips".[30]

Taxonomy

The following is a simplified classification of dinosaur groups based on their evolutionary relationships, and those of the
main dinosaur groups Theropoda, Sauropodomorpha and Ornithischia, compiled by Justin Tweet.[135] Further details
and other hypotheses of classification may be found on individual articles.

Dinosauria

†Ornithischia ("bird-hipped"; diverse bipedal and quadrupedal

herbivores)

†Heterodontosauridae (small herbivores/omnivores with

prominent canine-like teeth)
Restoration of six ornithopods; far
†Genasauria ("cheeked lizards")
left: Camptosaurus, left: Iguanodon,
center background:
†Thyreophora (armored dinosaurs; bipeds and
Shantungosaurus, center foreground:
quadrupeds)
Dryosaurus, right: Corythosaurus, far
right (large) Tenontosaurus.
†Eurypoda (heavy, quadrupedal
thyreophorans)

†Stegosauria (spikes and plates as primary armor)

†Huayangosauridae (small stegosaurs with flank osteoderms

and tail clubs)
†Stegosauridae (large stegosaurs)
 †Ankylosauria (scutes as primary armor)

†Parankylosauria (small, southern ankylosaurs with macuahuitl-

like tails)
†Nodosauridae (mostly spiky, club-less ankylosaurs)
†Ankylosauridae (characterized by flat scutes)

†Ankylosaurinae (club-tailed ankylosaurids)

†Neornithischia ("new ornithischians")

†Cerapoda ("horned feet")

†Marginocephalia (characterized by a cranial growth)

†Pachycephalosauria
(bipeds with domed or
knobby growth on skulls)
†Ceratopsia (bipeds and
quadrupeds; many had neck
frills and horns)

†Chaoyangsauridae
Restoration of four ceratopsids: top
(small, frill-less basal
ceratopsians) left - Triceratops, top right -
Styracosaurus, bottom left -
†Neoceratopsia ("new
Anchiceratops, bottom right -
ceratopsians")
Chasmosaurus.

†Leptoceratopsidae (little to no frills, hornless, with

robust jaws)
†Protoceratopsidae (basal ceratopsians with small
frills and stubby horns)

†Ceratopsoidea (large-horned ceratopsians)

†Ceratopsidae (large, elaborately

ornamented ceratopsians)

†Chasmosaurinae (ceratopsids with

enlarged brow horns)
†Centrosaurinae (ceratopsids
mostly characterized by frill and
nasal ornamentation)

†Nasutoceratopsini
(centrosaurines with enlarged
nasal cavities)
†Centrosaurini
(centrosaurines with enlarged
nasal horns)
†Pachyrhinosaurini (mostly
had nasal bosses instead of
horns)

†Ornithopoda (various sizes; bipeds and quadrupeds; evolved a

method of chewing using skull flexibility and numerous teeth)
†Jeholosauridae (small Asian neornithischians)
†Thescelosauridae ("wondrous lizards")

†Orodrominae (burrowers)
†Thescelosaurinae (large thescelosaurids)

†Iguanodontia ("iguana teeth"; advanced ornithopods)

†Elasmaria (mostly southern ornithopods with mineralized

plates along the ribs; may be thescelosaurids)
†Rhabdodontomorpha (with distinctive dentition)

†Rhabdodontidae (European rhabdodontomorphs)

†Dryosauridae (mid-sized, small headed)

†Ankylopollexia (early members mid-sized, stocky)

†Styracosterna ("spiked sterna")

†Hadrosauriformes (ancestrally had a

thumb spike; large quadrupedal
herbivores, with teeth merged into dental
batteries)

†Hadrosauromorpha (hadrosaurids
and their closest relatives)

†Hadrosauridae ("duck-billed
dinosaurs"; often with crests)

†Saurolophinae
(hadrosaurids with solid,
small, no crests)

†Brachylophosaurini
(short-crested)
†Kritosaurini
(enlarged, solid
nasal crests)
†Saurolophini
(small, spike-like
crests)
†Edmontosaurini
(flat-headed
saurolophines)

†Lambeosaurinae
(hadrosaurids often with
hollow crests)

†Aralosaurini
(solid-crested)
†Tsintaosaurini
(vertical, tube-like
crests)
†Parasaurolophini
(long, backwards-
 arcing crests)
†Lambeosaurini
(usually rounded
crests)

Saurischia

†Herrerasauridae (early bipedal carnivores)

†Sauropodomorpha (herbivores with small heads, long

necks, and long tails)

†Unaysauridae (primitive, strictly bipedal

"prosauropods")
†Plateosauria (diverse; bipeds and quadrupeds)

†Massospondylidae (long-necked, primitive Restoration of four macronarian

sauropodomorphs) sauropods: from left to right
†Riojasauridae (large, primitive Camarasaurus, Brachiosaurus,
sauropodomorphs) Giraffatitan, and Euhelopus

†Sauropodiformes (heavy, bipeds and

quadrupeds)

†Sauropoda (very large and heavy; quadrupedal)

†Lessemsauridae (gigantic yet lacking several weight-saving

adaptations)
†Gravisauria ("heavy lizards")

†Eusauropoda ("true sauropods")

†Turiasauria (often large, widespread sauropods)

†Neosauropoda ("new sauropods"; columnar limbs)

†Diplodocoidea (skulls and tails elongated;

teeth typically narrow and pencil-like)

†Rebbachisauridae (short-necked, low-

browsing diplodocoids often with high
backs)
†Flagellicaudata (whip-tailed)

†Dicraeosauridae (small, short-

necked diplodocoids with enlarged
cervical and dorsal vertebrae)
†Diplodocidae (extremely long-
necked)

†Apatosaurinae (robust
cervical vertebrae)
†Diplodocinae (long, thin
necks)

†Macronaria (boxy skulls; spoon- or pencil-

shaped teeth)

†Titanosauriformes ("titan lizard forms")

 †Brachiosauridae (long-necked,
long-armed macronarians)
†Somphospondyli ("porous
vertebrae")

†Euhelopodidae (stocky,
mostly Asian)
†Titanosauria (diverse; stocky,
with wide hips; most common
in the Late Cretaceous of
southern continents)

Theropoda (carnivorous)

Neotheropoda ("new theropods")

†Coelophysoidea (early theropods; includes Coelophysis and close relatives)

†"Dilophosaur-grade neotheropods" (larger kink-snouted dinosaurs)
Averostra ("bird snouts")

†Ceratosauria (generally elaborately horned carnivores that existed from the

Jurassic to Cretaceous periods, originally included Coelophysoidea)

†Ceratosauridae (ceratosaurs with large teeth)

†Abelisauroidea (ceratosaurs exemplified by reduced arms and
hands)

†Abelisauridae (large abelisauroids with short arms and

oftentimes elaborate facial ornamentation)
†Noasauridae (diverse, generally light theropods; may include
several obscure taxa)

†Elaphrosaurinae (bird-like; omnivorous as juveniles but

herbivorous as adults)
†Noasaurinae (small carnivores)

Tetanurae (stiff-tailed dinosaurs)

†Megalosauroidea (early group of large carnivores)

†Piatnitzkysauridae (small basal megalosauroids endemic to the

Americas)
†Megalosauridae (large megalosauroids with powerful arms and
hands)
†Spinosauridae (crocodile-like, semiaquatic carnivores)

Avetheropoda ("bird theropods")

†Megaraptora (theropods with large hand claws; either

carnosaurs or coelurosaurs, potentially tyrannosauroids)
†Carnosauria (large meat-eating dinosaurs; megalosauroids
sometimes included)

†Metriacanthosauridae (primitive Asian allosauroids)

†Allosauridae (Allosaurus and its very closest relatives)
†Carcharodontosauria (robust allosauroids)
 †Carcharodontosauridae (includes some of the
largest purely terrestrial carnivores)
†Neovenatoridae ("new hunters"; may include
megaraptorans)

Coelurosauria (feathered theropods, with a range of body sizes

and niches)

†"Nexus of basal coelurosaurs" (used by Tweet to denote

well-known taxa with unstable positions at the base of
Coelurosauria)
Tyrannoraptora ("tyrant thieves")

†Compsognathidae (small early coelurosaurs with

short forelimbs)
†Tyrannosauroidea (mostly large, primitive
coelurosaurs)

†Proceratosauridae (tyrannosauroids with head

crests)
†Tyrannosauridae (Tyrannosaurus and close
relatives)

Maniraptoriformes (bird-like dinosaurs)

†Ornithomimosauria (small-headed, mostly

toothless, omnivorous or possible herbivores)

†Ornithomimidae (very ostrich-like

dinosaurs)

Maniraptora (dinosaurs with pennaceous

feathers)

Restoration of six dromaeosaurid

theropods: from left to right
Microraptor, Velociraptor,
Austroraptor, Dromaeosaurus,
Utahraptor, and Deinonychus

†Alvarezsauroidea (small hunters with

reduced forelimbs)

†Alvarezsauridae (insectivores with

only one enlarged digit)

†Therizinosauria (tall, long-necked

theropods; omnivores and herbivores)

†Therizinosauroidea (larger
therizinosaurs)

†Therizinosauridae (sloth-like
herbivores, often with
 enlarged claws)

†Oviraptorosauria (omnivorous, beaked

dinosaurs)

†Caudipteridae (bird-like, basal

oviraptorosaurs)
†Caenagnathoidea (cassowary-like
oviraptorosaurs)

†Caenagnathidae (toothless
oviraptorosaurs known from
North America and Asia)
†Oviraptoridae (characterized
by two bony projections at the
back of the mouth; exclusive to
Asia)

Paraves (avialans and their closest

relatives)

†Scansoriopterygidae (small tree-

climbing theropods with
membranous wings)
†Deinonychosauria (toe-clawed
dinosaurs; may not form a natural
group)

†Archaeopterygidae (small,
winged theropods or primitive
birds)
†Troodontidae (omnivores;
enlarged brain cavities)
†Dromaeosauridae ("raptors")

†Microraptoria
(characterized by large
wings on both the arms
and legs; may have
been capable of
powered flight)
†Eudromaeosauria
(hunters with greatly
enlarged sickle claws)

†Unenlagiidae (piscivores;
may be dromaeosaurids)

†Halszkaraptorinae
(duck-like; potentially
semiaquatic)
†Unenlagiinae (long-
snouted)

Avialae (modern birds and extinct

relatives)

Timeline of major groups

Timeline of major dinosaur groups per Holtz (2007).

Paleobiology
Knowledge about dinosaurs is derived from a variety of fossil and non-fossil records, including fossilized bones, feces,
trackways, gastroliths, feathers, impressions of skin, internal organs and other soft tissues.[90][94] Many fields of study
contribute to our understanding of dinosaurs, including physics (especially biomechanics), chemistry, biology, and the
Earth sciences (of which paleontology is a sub-discipline).[136][137] Two topics of particular interest and study have been
dinosaur size and behavior.[138]

Size

Current evidence suggests that dinosaur average size varied through the Triassic, Early Jurassic, Late Jurassic and
Cretaceous.[111] Predatory theropod dinosaurs, which occupied most terrestrial carnivore niches during the Mesozoic,
most often fall into the 100 to 1 000 kg (220 to 2 200 lb) category when sorted by estimated weight into categories based
on order of magnitude, whereas recent predatory carnivoran mammals peak in the 10 to 100  kg (22 to 220  lb)
category.[139] The mode of Mesozoic dinosaur body masses is between 1 and 10 metric tons (1.1 and 11.0 short
tons).[140] This contrasts sharply with the average size of Cenozoic mammals, estimated by the National Museum of
Natural History as about 2 to 5 kg (4.4 to 11.0 lb).[141]

The sauropods were the largest and heaviest dinosaurs. For much of the dinosaur era, the smallest sauropods were larger
than anything else in their habitat, and the largest was an order of magnitude more massive than anything else that has
since walked the Earth. Giant prehistoric mammals such as Paraceratherium (the largest land mammal ever) were
dwarfed by the giant sauropods, and only modern whales
approach or surpass them in size.[142] There are several proposed
advantages for the large size of sauropods, including protection
from predation, reduction of energy use, and longevity, but it may
be that the most important advantage was dietary. Large animals
are more efficient at digestion than small animals, because food
spends more time in their digestive systems. This also permits Scale diagram comparing the average human to the
them to subsist on food with lower nutritive value than smaller longest known dinosaurs in five major clades:
animals. Sauropod remains are mostly found in rock formations    Sauropoda (Supersaurus vivianae)
interpreted as dry or seasonally dry, and the ability to eat large    Ornithopoda (Shantungosaurus giganteus)
quantities of low-nutrient browse would have been advantageous
   Theropoda (Spinosaurus aegyptiacus)
in such environments.[143]
   Thyreophora (Stegosaurus ungulatus)
   Marginocephalia (Triceratops prorsus)
Largest and smallest

Scientists will probably never be certain of the largest and smallest dinosaurs to have ever existed. This is because only a
tiny percentage of animals were ever fossilized and most of these remain buried in the earth. Few of the specimens that
are recovered are complete skeletons, and impressions of skin and other soft tissues are rare. Rebuilding a complete
skeleton by comparing the size and morphology of bones to those of similar, better-known species is an inexact art, and
reconstructing the muscles and other organs of the living animal is, at best, a process of educated guesswork.[144]

The tallest and heaviest dinosaur known from good skeletons is Giraffatitan
brancai (previously classified as a species of Brachiosaurus). Its remains were
discovered in Tanzania between 1907 and 1912. Bones from several similar-sized
individuals were incorporated into the skeleton now mounted and on display at
the Museum für Naturkunde in Berlin;[145] this mount is 12 meters (39 ft) tall and
Comparative size of Argentinosaurus 21.8 to 22.5 meters (72 to 74 ft) long,[146][147] and would have belonged to an
to the average human animal that weighed between 30 000 and 60 000  kilograms (70 000 and
130 000  lb). The longest complete dinosaur is the 27 meters (89  ft) long
Diplodocus, which was discovered in Wyoming in the United States and
displayed in Pittsburgh's Carnegie Museum of Natural History in 1907.[148] The longest dinosaur known from good
fossil material is Patagotitan: the skeleton mount in the American Museum of Natural History in New York is 37 meters
(121 ft) long. The Museo Municipal Carmen Funes in Plaza Huincul, Argentina, has an Argentinosaurus reconstructed
skeleton mount that is 39.7 meters (130 ft) long.[149]

There were larger dinosaurs, but knowledge of them is based entirely on a small
number of fragmentary fossils. Most of the largest herbivorous specimens on
record were discovered in the 1970s or later, and include the massive
Argentinosaurus, which may have weighed 80 000 to 100 000 kilograms (90 to
110 short tons) and reached lengths of 30 to 40 meters (98 to 131 ft); some of the
longest were the 33.5-meter (110  ft) long Diplodocus hallorum[143] (formerly
Seismosaurus), the 33-to-34-meter (108 to 112 ft) long Supersaurus,[150] and 37-
meter (121  ft) long Patagotitan; and the tallest, the 18-meter (59  ft) tall
Sauroposeidon, which could have reached a sixth-floor window. The heaviest
and longest dinosaur may have been Maraapunisaurus, known only from a now
lost partial vertebral neural arch described in 1878. Extrapolating from the
illustration of this bone, the animal may have been 58 meters (190  ft) long and
weighed 122 400 kg (270 000 lb).[143] However, as no further evidence of An adult bee hummingbird, the
sauropods of this size has been found, and the discoverer, Cope, had made smallest known dinosaur
typographic errors before, it is likely to have been an extreme overestimation.[151]

The largest carnivorous dinosaur was Spinosaurus, reaching a length of 12.6 to 18 meters (41 to 59 ft), and weighing 7
to 20.9 metric tons (7.7 to 23.0 short tons).[152][153] Other large carnivorous theropods included Giganotosaurus,
Carcharodontosaurus and Tyrannosaurus.[153] Therizinosaurus and Deinocheirus were among the tallest of the
theropods. The largest ornithischian dinosaur was probably the hadrosaurid Shantungosaurus giganteus which measured
16.6 meters (54 ft).[154] The largest individuals may have weighed as much as 16 metric tons (18 short tons).[155]
The smallest dinosaur known is the bee hummingbird,[156] with a length of only 5 centimeters (2.0  in) and mass of
around 1.8 g (0.063 oz).[157] The smallest known non-avialan dinosaurs were about the size of pigeons and were those
theropods most closely related to birds.[158] For example, Anchiornis huxleyi is currently the smallest non-avialan
dinosaur described from an adult specimen, with an estimated weight of 110 g (3.9 oz)[159] and a total skeletal length of
34 centimeters (1.12  ft).[158][159] The smallest herbivorous non-avialan dinosaurs included Microceratus and
Wannanosaurus, at about 60 centimeters (2.0 ft) long each.[160][161]

Behavior

Many modern birds are highly social, often found living in flocks. There is
general agreement that some behaviors that are common in birds, as well as in
crocodiles (closest living relatives of birds), were also common among extinct
dinosaur groups. Interpretations of behavior in fossil species are generally based
on the pose of skeletons and their habitat, computer simulations of their
biomechanics, and comparisons with modern animals in similar ecological
niches.[136]

The first potential evidence for herding or flocking as a widespread behavior

A nesting ground of the hadrosaur
Maiasaura peeblesorum was
common to many dinosaur groups in addition to birds was the 1878 discovery of
31 Iguanodon, ornithischians that were then thought to have perished together in
discovered in 1978
Bernissart, Belgium, after they fell into a deep, flooded sinkhole and
drowned.[162] Other mass-death sites have been discovered subsequently. Those,
along with multiple trackways, suggest that gregarious behavior was common in many early dinosaur species. Trackways
of hundreds or even thousands of herbivores indicate that duck-billed (hadrosaurids) may have moved in great herds, like
the American bison or the African springbok. Sauropod tracks document that these animals traveled in groups composed
of several different species, at least in Oxfordshire, England,[163] although there is no evidence for specific herd
structures.[164] Congregating into herds may have evolved for defense, for migratory purposes, or to provide protection
for young. There is evidence that many types of slow-growing dinosaurs, including various theropods, sauropods,
ankylosaurians, ornithopods, and ceratopsians, formed aggregations of immature individuals. One example is a site in
Inner Mongolia that has yielded remains of over 20 Sinornithomimus, from one to seven years old. This assemblage is
interpreted as a social group that was trapped in mud.[165] The interpretation of dinosaurs as gregarious has also extended
to depicting carnivorous theropods as pack hunters working together to bring down large prey.[166][167] However, this
lifestyle is uncommon among modern birds, crocodiles, and other reptiles, and the taphonomic evidence suggesting
mammal-like pack hunting in such theropods as Deinonychus and Allosaurus can also be interpreted as the results of fatal
disputes between feeding animals, as is seen in many modern diapsid predators.[168]

The crests and frills of some dinosaurs, like the marginocephalians, theropods and
lambeosaurines, may have been too fragile to be used for active defense, and so
they were likely used for sexual or aggressive displays, though little is known
about dinosaur mating and territorialism. Head wounds from bites suggest that
theropods, at least, engaged in active aggressive confrontations.[169]

From a behavioral standpoint, one of the most valuable dinosaur fossils was Restoration of two Centrosaurus
discovered in the Gobi Desert in 1971. It included a Velociraptor attacking a apertus engaged in intra-specific
Protoceratops,[170] providing evidence that dinosaurs did indeed attack each combat
other. [171] Additional evidence for attacking live prey is the partially healed tail of
an Edmontosaurus, a hadrosaurid dinosaur; the tail is damaged in such a way that
shows the animal was bitten by a tyrannosaur but survived.[171] Cannibalism amongst some species of dinosaurs was
confirmed by tooth marks found in Madagascar in 2003, involving the theropod Majungasaurus.[172]

Comparisons between the scleral rings of dinosaurs and modern birds and reptiles have been used to infer daily activity
patterns of dinosaurs. Although it has been suggested that most dinosaurs were active during the day, these comparisons
have shown that small predatory dinosaurs such as dromaeosaurids, Juravenator, and Megapnosaurus were likely
nocturnal. Large and medium-sized herbivorous and omnivorous dinosaurs such as ceratopsians, sauropodomorphs,
hadrosaurids, ornithomimosaurs may have been cathemeral, active during short intervals throughout the day, although the
small ornithischian Agilisaurus was inferred to be diurnal.[173]
Based on fossil evidence from dinosaurs such as Oryctodromeus, some ornithischian species seem to have led a partially
fossorial (burrowing) lifestyle.[174] Many modern birds are arboreal (tree climbing), and this was also true of many
Mesozoic birds, especially the enantiornithines.[175] While some early bird-like species may have already been arboreal
as well (including dromaeosaurids) such as Microraptor[176]) most non-avialan dinosaurs seem to have relied on land-
based locomotion. A good understanding of how dinosaurs moved on the ground is key to models of dinosaur behavior;
the science of biomechanics, pioneered by Robert McNeill Alexander, has provided significant insight in this area. For
example, studies of the forces exerted by muscles and gravity on dinosaurs' skeletal structure have investigated how fast
dinosaurs could run,[136] whether diplodocids could create sonic booms via whip-like tail snapping,[177] and whether
sauropods could float.[178]

Communication

Modern birds are known to communicate using visual and auditory signals, and the wide diversity of visual display
structures among fossil dinosaur groups, such as horns, frills, crests, sails, and feathers, suggests that visual
communication has always been important in dinosaur biology.[179] Reconstruction of the plumage color of Anchiornis,
suggest the importance of color in visual communication in non-avian dinosaurs.[180] Vocalization in non-avian
dinosaurs is less certain. In birds, the larynx plays no role in sound production. Instead they vocalize with a novel organ
called the syrinx, located further down the trachea.[181] The earliest remains of a syrinx was found in a specimen of the
duck-like Vegavis iaai dated 69 –66  million years ago, and this organ is unlikely to have existed in non-avian
dinosaurs.[182]

Paleontologist Phil Senter has suggested that since non-avian dinosaurs did not
have a syrinx, and their next closest living relatives, crocodilians, use the larynx,
they could not vocalize as the common ancestor would have been mute. He states
that they mostly on visual displays and possibly non-vocal acoustic sounds like
hissing, jaw grinding or clapping, splashing and wing beating (possible in winged
maniraptoran dinosaurs).[179] Other researchers have countered that vocalizations
also exist in turtles, the closest relatives of archosaurs, suggesting that the trait is
ancestral to their lineage. In addition, vocal communication in dinosaurs is
indicated by the development of advanced hearing in nearly all major groups.
Hence the syrinx may have supplemented and then replaced the larynx as a vocal
organ rather than there being a "silent period" in bird evolution.[183]
Restoration of a striking and unusual
In 2023, a fossilized larynx was described from a specimen of the ankylosaurid visual display in a Lambeosaurus
Pinacosaurus. The structure was composed of cricoid and arytenoid cartilages, magnicristatus. The crest could also
similar to those of non-avian reptiles. However, the mobile cricoid-arytenoid joint have acted as a resonating chamber
and long arytenoid cartilages would have allowed for air-flow control similar to for sounds
that of birds, and thus could have made bird-like vocalizations. In addition, the
cartilages were ossified, implying that laryngeal ossification is a feature of some
non-avian dinosaurs.[184] A 2016 study concludes that some dinosaurs may have produced closed mouth vocalizations
like cooing, hooting and booming. These occur in both reptiles and birds and involve inflating the esophagus or tracheal
pouches. Such vocalizations evolved independently in extant archosaurs numerous times, following increases in body
size.[185] The crests of some hadrosaurids and the nasal chambers of ankylosaurids have been suggested to have
functioned in acoustic resonance.[186][187]

Reproductive biology

All dinosaurs laid amniotic eggs. Dinosaur eggs were usually laid in a nest. Most species create somewhat elaborate nests
which can be cups, domes, plates, beds scrapes, mounds, or burrows.[188] Some species of modern bird have no nests;
the cliff-nesting common guillemot lays its eggs on bare rock, and male emperor penguins keep eggs between their body
and feet. Primitive birds and many non-avialan dinosaurs often lay eggs in communal nests, with males primarily
incubating the eggs. While modern birds have only one functional oviduct and lay one egg at a time, more primitive birds
and dinosaurs had two oviducts, like crocodiles. Some non-avialan dinosaurs, such as Troodon, exhibited iterative laying,
where the adult might lay a pair of eggs every one or two days, and then ensured simultaneous hatching by delaying
brooding until all eggs were laid.[189]
 When laying eggs, females grow a special type of bone between the hard outer
bone and the marrow of their limbs. This medullary bone, which is rich in
calcium, is used to make eggshells. A discovery of features in a Tyrannosaurus
skeleton provided evidence of medullary bone in extinct dinosaurs and, for the
first time, allowed paleontologists to establish the sex of a fossil dinosaur
specimen. Further research has found medullary bone in the carnosaur Allosaurus
and the ornithopod Tenontosaurus. Because the line of dinosaurs that includes
Allosaurus and Tyrannosaurus diverged from the line that led to Tenontosaurus
very early in the evolution of dinosaurs, this suggests that the production of
Nest of a plover (Charadrius) medullary tissue is a general characteristic of all dinosaurs.[190]

Another widespread trait among modern

birds (but see below in regards to fossil groups and extant megapodes) is parental
care for young after hatching. Jack Horner's 1978 discovery of a Maiasaura
("good mother lizard") nesting ground in Montana demonstrated that parental
care continued long after birth among ornithopods.[191] A specimen of the
oviraptorid Citipati osmolskae was discovered in a chicken-like brooding position
in 1993,[192] which may indicate that they had begun using an insulating layer of
feathers to keep the eggs warm.[193] An embryo of the basal sauropodomorph
Massospondylus was found without teeth, indicating that some parental care was Fossil interpreted as a nesting
required to feed the young dinosaurs.[194] Trackways have also confirmed oviraptorid Citipati at the American
parental behavior among ornithopods from the Isle of Skye in northwestern Museum of Natural History. Smaller
Scotland.[195] fossil far right showing inside one of
the eggs.
However, there is ample evidence of precociality or superprecociality among
many dinosaur species, particularly theropods. For instance, non-ornithuromorph
birds have been abundantly demonstrated to have had slow growth rates, megapode-like egg burying behavior and the
ability to fly soon after birth.[196][197][198][199] Both Tyrannosaurus and Troodon had juveniles with clear
superprecociality and likely occupying different ecological niches than the adults.[189] Superprecociality has been
inferred for sauropods.[200]

Genital structures are unlikely to fossilize as they lack scales that may allow preservation via pigmentation or residual
calcium phosphate salts. In 2021, the best preserved specimen of a dinosaur's cloacal vent exterior was described for
Psittacosaurus, demonstrating lateral swellings similar to crocodylian musk glands used in social displays by both sexes
and pigmented regions which could also reflect a signalling function. However, this specimen on its own does not offer
enough information to determine whether this dinosaur had sexual signalling functions; it only supports the possibility.
Cloacal visual signalling can occur in either males or females in living birds, making it unlikely to be useful to determine
sex for extinct dinosaurs.[201]

Physiology

Because both modern crocodilians and birds have four-chambered hearts (albeit modified in crocodilians), it is likely that
this is a trait shared by all archosaurs, including all dinosaurs.[202] While all modern birds have high metabolisms and are
endothermic ("warm-blooded"), a vigorous debate has been ongoing since the 1960s regarding how far back in the
dinosaur lineage this trait extended. Various researchers have supported dinosaurs as being endothermic, ectothermic
("cold-blooded"), or somewhere in between.[203] An emerging consensus among researchers is that, while different
lineages of dinosaurs would have had different metabolisms, most of them had higher metabolic rates than other reptiles
but lower than living birds and mammals,[204] which is termed mesothermy by some.[205] Evidence from crocodiles and
their extinct relatives suggests that such elevated metabolisms could have developed in the earliest archosaurs, which
were the common ancestors of dinosaurs and crocodiles.[206][207]

After non-avian dinosaurs were discovered, paleontologists first posited that they were ectothermic. This was used to
imply that the ancient dinosaurs were relatively slow, sluggish organisms, even though many modern reptiles are fast and
light-footed despite relying on external sources of heat to regulate their body temperature. The idea of dinosaurs as
ectothermic remained a prevalent view until Robert T. Bakker, an early proponent of dinosaur endothermy, published an
influential paper on the topic in 1968. Bakker specifically used anatomical and ecological evidence to argue that
sauropods, which had hitherto been depicted as sprawling aquatic animals with their tails dragging on the ground, were
 endotherms that lived vigorous, terrestrial lives. In 1972, Bakker expanded on his
arguments based on energy requirements and predator-prey ratios. This was one
of the seminal results that led to the dinosaur renaissance.[63][64][60][208]

One of the greatest contributions to the modern understanding of dinosaur

physiology has been paleohistology, the study of microscopic tissue structure in
dinosaurs.[209][210] From the 1960s forward, Armand de Ricqlès suggested that
the presence of fibrolamellar bone—bony tissue with an irregular, fibrous texture
This 1897 restoration of and filled with blood vessels—was indicative of consistently fast growth and
Brontosaurus as an aquatic, tail- therefore endothermy. Fibrolamellar bone was common in both dinosaurs and
dragging animal, by Charles R. pterosaurs,[211][212] though not universally present.[213][214] This has led to a
Knight, typified early views on significant body of work in reconstructing growth curves and modeling the
dinosaur lifestyles. evolution of growth rates across various dinosaur lineages,[215] which has
suggested overall that dinosaurs grew faster than living reptiles.[210] Other lines
of evidence suggesting endothermy include the presence of feathers and other
types of body coverings in many lineages (see § Feathers); more consistent ratios of the isotope oxygen-18 in bony tissue
compared to ectotherms, particularly as latitude and thus air temperature varied, which suggests stable internal
temperatures[216][217] (although these ratios can be altered during fossilization[218]); and the discovery of polar
dinosaurs, which lived in Australia, Antarctica, and Alaska when these places would have had cool, temperate
climates.[219][220][221][222]

In saurischian dinosaurs, higher metabolisms were supported by the evolution of

the avian respiratory system, characterized by an extensive system of air sacs that
extended the lungs and invaded many of the bones in the skeleton, making them
hollow.[223] Such respiratory systems, which may have appeared in the earliest
saurischians,[224] would have provided them with more oxygen compared to a
mammal of similar size, while also having a larger resting tidal volume and
requiring a lower breathing frequency, which would have allowed them to sustain
higher activity levels.[142] The rapid airflow would also have been an effective
Comparison between the air sacs of
cooling mechanism, which in conjunction with a lower metabolic rate[225] would
an abelisaur and a bird
have prevented large sauropods from overheating. These traits may have enabled
sauropods to grow quickly to gigantic sizes.[226][227] Sauropods may also have
benefitted from their size—their small surface area to volume ratio meant that they would have been able to
thermoregulate more easily, a phenomenon termed gigantothermy.[142][228]

Like other reptiles, dinosaurs are primarily uricotelic, that is, their kidneys extract nitrogenous wastes from their
bloodstream and excrete it as uric acid instead of urea or ammonia via the ureters into the intestine. This would have
helped them to conserve water.[204] In most living species, uric acid is excreted along with feces as a semisolid
waste.[229][230] However, at least some modern birds (such as hummingbirds) can be facultatively ammonotelic,
excreting most of the nitrogenous wastes as ammonia.[231] This material, as well as the output of the intestines, emerges
from the cloaca.[232][233] In addition, many species regurgitate pellets,[234] and fossil pellets are known as early as the
Jurassic from Anchiornis.[235]

The size and shape of the brain can be partly reconstructed based on the surrounding bones. In 1896, Marsh calculated
ratios between brain weight and body weight of seven species of dinosaurs, showing that the brain of dinosaurs was
proportionally smaller than in today's crocodiles, and that the brain of Stegosaurus was smaller than in any living land
vertebrate. This contributed to the widespread public notion of dinosaurs as being sluggish and extraordinarily stupid.
Harry Jerison, in 1973, showed that proportionally smaller brains are expected at larger body sizes, and that brain size in
dinosaurs was not smaller than expected when compared to living reptiles.[236] Later research showed that relative brain
size progressively increased during the evolution of theropods, with the highest intelligence – comparable to that of
modern birds – calculated for the troodontid Troodon.[237]

Origin of birds
The possibility that dinosaurs were the ancestors of birds was first suggested in 1868 by Thomas Henry Huxley.[238]
After the work of Gerhard Heilmann in the early 20th century, the theory of birds as dinosaur descendants was
abandoned in favor of the idea of them being descendants of generalized thecodonts, with the key piece of evidence
being the supposed lack of clavicles in dinosaurs.[239] However, as later discoveries showed, clavicles (or a single fused
wishbone, which derived from separate clavicles) were not actually absent;[14] they had been found as early as 1924 in
Oviraptor, but misidentified as an interclavicle.[240] In the 1970s, Ostrom revived the dinosaur–bird theory,[241] which
gained momentum in the coming decades with the advent of cladistic analysis,[242] and a great increase in the discovery
of small theropods and early birds.[32] Of particular note have been the fossils of the Jehol Biota, where a variety of
theropods and early birds have been found, often with feathers of some type.[70][14] Birds share over a hundred distinct
anatomical features with theropod dinosaurs, which are now generally accepted to have been their closest ancient
relatives.[243] They are most closely allied with maniraptoran coelurosaurs.[14] A minority of scientists, most notably
Alan Feduccia and Larry Martin, have proposed other evolutionary paths, including revised versions of Heilmann's basal
archosaur proposal,[244] or that maniraptoran theropods are the ancestors of birds but themselves are not dinosaurs, only
convergent with dinosaurs.[245]

Feathers

Feathers are one of the most recognizable characteristics of modern birds, and a
trait that was also shared by several non-avian dinosaurs. Based on the current
distribution of fossil evidence, it appears that feathers were an ancestral
dinosaurian trait, though one that may have been selectively lost in some
species.[246] Direct fossil evidence of feathers or feather-like structures has been
discovered in a diverse array of species in many non-avian dinosaur groups,[70]
both among saurischians and ornithischians. Simple, branched, feather-like
structures are known from heterodontosaurids, primitive neornithischians,[247]
and theropods,[248] and primitive ceratopsians. Evidence for true, vaned feathers
similar to the flight feathers of modern birds has been found only in the theropod
subgroup Maniraptora, which includes oviraptorosaurs, troodontids,
dromaeosaurids, and birds.[14][249] Feather-like structures known as pycnofibres Various feathered non-avian
have also been found in pterosaurs.[250] dinosaurs, including Archaeopteryx,
Anchiornis, Microraptor and
However, researchers do not agree regarding whether these structures share a Zhenyuanlong
common origin between lineages (i.e., they are homologous),[251][252] or if they
were the result of widespread experimentation with skin coverings among
ornithodirans.[253] If the former is the case, filaments may have been common in the ornithodiran lineage and evolved
before the appearance of dinosaurs themselves.[246] Research into the genetics of American alligators has revealed that
crocodylian scutes do possess feather-keratins during embryonic development, but these keratins are not expressed by the
animals before hatching.[254] The description of feathered dinosaurs has not been without controversy in general;
perhaps the most vocal critics have been Alan Feduccia and Theagarten Lingham-Soliar, who have proposed that some
purported feather-like fossils are the result of the decomposition of collagenous fiber that underlaid the dinosaurs'
skin,[255][256][257] and that maniraptoran dinosaurs with vaned feathers were not actually dinosaurs, but convergent with
dinosaurs.[245][256] However, their views have for the most part not been accepted by other researchers, to the point that
the scientific nature of Feduccia's proposals has been questioned.[258]

Archaeopteryx was the first fossil found that revealed a potential connection between dinosaurs and birds. It is considered
a transitional fossil, in that it displays features of both groups. Brought to light just two years after Charles Darwin's
seminal On the Origin of Species (1859), its discovery spurred the nascent debate between proponents of evolutionary
biology and creationism. This early bird is so dinosaur-like that, without a clear impression of feathers in the surrounding
rock, at least one specimen was mistaken for the small theropod Compsognathus.[259] Since the 1990s, a number of
additional feathered dinosaurs have been found, providing even stronger evidence of the close relationship between
dinosaurs and modern birds. Many of these specimens were unearthed in the lagerstätten of the Jehol Biota.[252] If
feather-like structures were indeed widely present among non-avian dinosaurs, the lack of abundant fossil evidence for
them may be due to the fact that delicate features like skin and feathers are seldom preserved by fossilization and thus
often absent from the fossil record.[260]

Skeleton
Because feathers are often associated with birds, feathered dinosaurs are often touted as the missing link between birds
and dinosaurs. However, the multiple skeletal features also shared by the two groups represent another important line of
evidence for paleontologists. Areas of the skeleton with important similarities include the neck, pubis, wrist (semi-lunate
carpal), arm and pectoral girdle, furcula (wishbone), and breast bone. Comparison of bird and dinosaur skeletons through
cladistic analysis strengthens the case for the link.[261]

Soft anatomy

Large meat-eating dinosaurs had a complex system of air sacs similar to those
found in modern birds, according to a 2005 investigation led by Patrick M.
O'Connor. The lungs of theropod dinosaurs (carnivores that walked on two legs
and had bird-like feet) likely pumped air into hollow sacs in their skeletons, as is
the case in birds. "What was once formally considered unique to birds was
present in some form in the ancestors of birds", O'Connor said.[262][263] In 2008,
scientists described Aerosteon riocoloradensis, the skeleton of which supplies the
strongest evidence to date of a dinosaur with a bird-like breathing system. CT
scanning of Aerosteon's fossil bones revealed evidence for the existence of air
sacs within the animal's body cavity.[223][264]

Behavioral evidence

Fossils of the troodonts Mei and Sinornithoides demonstrate that some dinosaurs Pneumatopores on the left ilium of
slept with their heads tucked under their arms.[265] This behavior, which may Aerosteon riocoloradensis
have helped to keep the head warm, is also characteristic of modern birds.
Several deinonychosaur and oviraptorosaur specimens have also been found
preserved on top of their nests, likely brooding in a bird-like manner.[266] The ratio between egg volume and body mass
of adults among these dinosaurs suggest that the eggs were primarily brooded by the male, and that the young were
highly precocial, similar to many modern ground-dwelling birds.[267]

Some dinosaurs are known to have used gizzard stones like modern birds. These stones are swallowed by animals to aid
digestion and break down food and hard fibers once they enter the stomach. When found in association with fossils,
gizzard stones are called gastroliths.[268]

Extinction of major groups

All non-avian dinosaurs and most lineages of birds[269] became extinct in a mass extinction event, called the Cretaceous–
Paleogene (K-Pg) extinction event, at the end of the Cretaceous period. Above the Cretaceous–Paleogene boundary,
which has been dated to 66.038 ± 0.025 million years ago,[270] fossils of non-avian dinosaurs disappear abruptly; the
absence of dinosaur fossils was historically used to assign rocks to the ensuing Cenozoic. The nature of the event that
caused this mass extinction has been extensively studied since the 1970s, leading to the development of two mechanisms
that are thought to have played major roles: an extraterrestrial impact event in the Yucatán Peninsula, along with flood
basalt volcanism in India. However, the specific mechanisms of the extinction event and the extent of its effects on
dinosaurs are still areas of ongoing research.[271] Alongside dinosaurs, many other groups of animals became extinct:
pterosaurs, marine reptiles such as mosasaurs and plesiosaurs, several groups of mammals, ammonites (nautilus-like
mollusks), rudists (reef-building bivalves), and various groups of marine plankton.[272][273] In all, approximately 47% of
genera and 76% of species on Earth became extinct during the K-Pg extinction event.[274] The relatively large size of
most dinosaurs and the low diversity of small-bodied dinosaur species at the end of the Cretaceous may have contributed
to their extinction;[275] the extinction of the bird lineages that did not survive may also have been caused by a
dependence on forest habitats or a lack of adaptations to eating seeds for survival.[276][277]

Pre-extinction diversity
Just before the K-Pg extinction event, the number of non-avian dinosaur species that existed globally has been estimated
at between 628 and 1078.[278] It remains uncertain whether the diversity of dinosaurs was in gradual decline before the
K-Pg extinction event, or whether dinosaurs were actually thriving prior to the extinction. Rock formations from the
Maastrichtian epoch, which directly preceded the extinction, have been found to have lower diversity than the preceding
Campanian epoch, which led to the prevailing view of a long-term decline in diversity.[272][273][279] However, these
comparisons did not account either for varying preservation potential between rock units or for different extents of
exploration and excavation.[271] In 1984, Dale Russell carried out an analysis to account for these biases, and found no
evidence of a decline;[280] another analysis by David Fastovsky and colleagues in 2004 even showed that dinosaur
diversity continually increased until the extinction,[281] but this analysis has been rebutted.[282] Since then, different
approaches based on statistics and mathematical models have variously supported either a sudden
extinction[271][278][283] or a gradual decline.[284][285] End-Cretaceous trends in diversity may have varied between
dinosaur lineages: it has been suggested that sauropods were not in decline, while ornithischians and theropods were in
decline.[286][287]

Impact event

The bolide impact hypothesis, first brought to wide attention in 1980 by Walter Alvarez,
Luis Alvarez, and colleagues, attributes the K-Pg extinction event to a bolide
(extraterrestrial projectile) impact.[288] Alvarez and colleagues proposed that a sudden
increase in iridium levels, recorded around the world in rock deposits at the Cretaceous–
Paleogene boundary, was direct evidence of the impact.[289] Shocked quartz, indicative
of a strong shockwave emanating from an impact, was also found worldwide.[290] The
actual impact site remained elusive until a crater measuring 180  km (110  mi) wide was
discovered in the Yucatán Peninsula of southeastern Mexico, and was publicized in a
1991 paper by Alan Hildebrand and colleagues.[291] Now, the bulk of the evidence
suggests that a bolide 5 to 15 kilometers (3 to 91 ⁄2 miles) wide impacted the Yucatán
Peninsula 66 million years ago, forming this crater[292] and creating a "kill mechanism"
that triggered the extinction event.[293][294][295] Luis (left) and his son
Walter Alvarez (right) at the
Within hours, the Chicxulub impact would have created immediate effects such as K-T Boundary in Gubbio,
earthquakes,[296] tsunamis,[297] and a global firestorm that likely killed unsheltered Italy, 1981
animals and started wildfires.[298][299] However, it would also have had longer-term
consequences for the environment. Within days, sulfate aerosols released from
rocks at the impact site would have contributed to acid rain and ocean
acidification.[300][301] Soot aerosols are thought to have spread around the world
over the ensuing months and years; they would have cooled the surface of the
Earth by reflecting thermal radiation, and greatly slowed photosynthesis by
blocking out sunlight, thus creating an impact winter.[271][302][303] (This role was
ascribed to sulfate aerosols until experiments demonstrated otherwise.[301]) The
cessation of photosynthesis would have led to the collapse of food webs
depending on leafy plants, which included all dinosaurs save for grain-eating
birds.[277]

Deccan Traps

At the time of the K-Pg extinction, the Deccan Traps flood basalts of India were The Chicxulub Crater at the tip of the
actively erupting. The eruptions can be separated into three phases around the K- Yucatán Peninsula; the impactor that
Pg boundary, two prior to the boundary and one after. The second phase, which formed this crater may have caused
occurred very close to the boundary, would have extruded 70 to 80% of the the dinosaur extinction.
volume of these eruptions in intermittent pulses that occurred around 100,000
years apart.[304][305] Greenhouse gases such as carbon dioxide and sulfur dioxide
would have been released by this volcanic activity,[306][307] resulting in climate change through temperature
perturbations of roughly 3 °C (5.4 °F) but possibly as high as 7 °C (13 °F).[308] Like the Chicxulub impact, the eruptions
may also have released sulfate aerosols, which would have caused acid rain and global cooling.[309] However, due to
large error margins in the dating of the eruptions, the role of the Deccan Traps in the K-Pg extinction remains
unclear.[270][271][310]

Before 2000, arguments that the Deccan Traps eruptions—as opposed to the Chicxulub impact—caused the extinction
were usually linked to the view that the extinction was gradual. Prior to the discovery of the Chicxulub crater, the Deccan
Traps were used to explain the global iridium layer;[306][311] even after the crater's discovery, the impact was still thought
to only have had a regional, not global, effect on the extinction event.[312] In response, Luis Alvarez rejected volcanic
activity as an explanation for the iridium layer and the extinction as a whole.[313] Since then, however, most researchers
have adopted a more moderate position, which identifies the Chicxulub impact as the primary progenitor of the extinction
while also recognizing that the Deccan Traps may also have played a role. Walter Alvarez himself has acknowledged that
the Deccan Traps and other ecological factors may have contributed to the extinctions in addition to the Chicxulub
impact.[314] Some estimates have placed the start of the second phase in the Deccan Traps eruptions within 50,000 years
after the Chicxulub impact.[315] Combined with mathematical modelling of the seismic waves that would have been
generated by the impact, this has led to the suggestion that the Chicxulub impact may have triggered these eruptions by
increasing the permeability of the mantle plume underlying the Deccan Traps.[316][317]

Whether the Deccan Traps were a major cause of the extinction, on par with the Chicxulub impact, remains uncertain.
Proponents consider the climatic impact of the sulfur dioxide released to have been on par with the Chicxulub impact,
and also note the role of flood basalt volcanism in other mass extinctions like the Permian-Triassic extinction
event.[318][319] They consider the Chicxulub impact to have worsened the ongoing climate change caused by the
eruptions.[320] Meanwhile, detractors point out the sudden nature of the extinction and that other pulses in Deccan Traps
activity of comparable magnitude did not appear to have caused extinctions. They also contend that the causes of
different mass extinctions should be assessed separately.[321] In 2020, Alfio Chiarenza and colleagues suggested that the
Deccan Traps may even have had the opposite effect: they suggested that the long-term warming caused by its carbon
dioxide emissions may have dampened the impact winter from the Chicxulub impact.[295]

Possible Paleocene survivors

Non-avian dinosaur remains have occasionally been found above the K-Pg boundary. In 2000, Spencer Lucas and
colleagues reported the discovery of a single hadrosaur right femur in the San Juan Basin of New Mexico, and described
it as evidence of Paleocene dinosaurs. The rock unit in which the bone was discovered has been dated to the early
Paleocene epoch, approximately 64.8 million years ago.[322] If the bone was not re-deposited by weathering action, it
would provide evidence that some dinosaur populations survived at least half a million years into the Cenozoic.[323]
Other evidence includes the presence of dinosaur remains in the Hell Creek Formation up to 1.3 m (4.3 ft) above the
Cretaceous–Paleogene boundary, representing 40,000 years of elapsed time. This has been used to support the view that
the K-Pg extinction was gradual.[324] However, these supposed Paleocene dinosaurs are considered by many other
researchers to be reworked, that is, washed out of their original locations and then re-buried in younger
sediments.[325][326][327] The age estimates have also been considered unreliable.[328]

Cultural depictions
By human standards, dinosaurs were creatures of fantastic appearance and often
enormous size. As such, they have captured the popular imagination and become
an enduring part of human culture. The entry of the word "dinosaur" into the
common vernacular reflects the animals' cultural importance: in English,
"dinosaur" is commonly used to describe anything that is impractically large,
obsolete, or bound for extinction.[329]

Public enthusiasm for dinosaurs first developed in Victorian England, where in

1854, three decades after the first scientific descriptions of dinosaur remains, a Outdated Iguanodon statues created
menagerie of lifelike dinosaur sculptures was unveiled in London's Crystal Palace by Benjamin Waterhouse Hawkins
Park. The Crystal Palace dinosaurs proved so popular that a strong market in for the Crystal Palace Park in 1853
smaller replicas soon developed. In subsequent decades, dinosaur exhibits opened
at parks and museums around the world, ensuring that successive generations
would be introduced to the animals in an immersive and exciting way.[330] The enduring popularity of dinosaurs, in its
turn, has resulted in significant public funding for dinosaur science, and has
frequently spurred new discoveries. In the United States, for example, the
competition between museums for public attention led directly to the Bone Wars
of the 1880s and 1890s, during which a pair of feuding paleontologists made
enormous scientific contributions.[331]

The popular preoccupation with dinosaurs has ensured their appearance in

Gertie the Dinosaur (1914) by Winsor
literature, film, and other media. Beginning in 1852 with a passing mention in
McCay, featuring the first animated
Charles Dickens' Bleak House,[332] dinosaurs have been featured in large dinosaur
numbers of fictional works. Jules Verne's 1864 novel Journey to the Center of the
Earth, Sir Arthur Conan Doyle's 1912 book The Lost World, the 1914 animated
film Gertie the Dinosaur (featuring the first animated dinosaur), the iconic 1933 film King Kong, the 1954 Godzilla and
its many sequels, the best-selling 1990 novel Jurassic Park by Michael Crichton and its 1993 film adaptation are just a
few notable examples of dinosaur appearances in fiction. Authors of general-interest non-fiction works about dinosaurs,
including some prominent paleontologists, have often sought to use the animals as a way to educate readers about science
in general. Dinosaurs are ubiquitous in advertising; numerous companies have referenced dinosaurs in printed or
televised advertisements, either in order to sell their own products or in order to characterize their rivals as slow-moving,
dim-witted, or obsolete.[333][334]

See also
Dinosaur diet and feeding
Evolutionary history of life
Lists of dinosaur-bearing stratigraphic units
List of dinosaur genera
List of bird genera
List of birds
List of informally named dinosaurs
List of films featuring dinosaurs

Further reading
University of Southampton (September 29, 2021). "Two New Species of Large Predatory Dinosaur With
Crocodile-Like Skulls Discovered on Isle of Wight" (https://scitechdaily.com/two-new-species-of-large-pr
edatory-dinosaur-with-crocodile-like-skulls-discovered-on-isle-of-wight/). SciTechDaily.
Zhou, Zhonghe (October 2004). "The origin and early evolution of birds: discoveries, disputes, and
perspectives from fossil evidence" (https://web.archive.org/web/20110721144552/http://www.cisneros-he
redia.org/infotrans/usfq/ornitofauna/pdfs/zhou2004.pdf) (PDF). Naturwissenschaften. Berlin: Springer
Science+Business Media. 91 (10): 455–471. Bibcode:2004NW.....91..455Z (https://ui.adsabs.harvard.ed
u/abs/2004NW.....91..455Z). doi:10.1007/s00114-004-0570-4 (https://doi.org/10.1007%2Fs00114-004-05
70-4). ISSN 0028-1042 (https://www.worldcat.org/issn/0028-1042). PMID 15365634 (https://pubmed.ncb
i.nlm.nih.gov/15365634). S2CID 3329625 (https://api.semanticscholar.org/CorpusID:3329625). Archived
from the original (http://www.cisneros-heredia.org/infotrans/usfq/ornitofauna/pdfs/zhou2004.pdf) (PDF)
on July 21, 2011. Retrieved November 6, 2019.
Paul, Gregory S. (2002). Dinosaurs of the Air: The Evolution and Loss of Flight in Dinosaurs and Birds (h
ttps://archive.org/details/dinosaursofairev0000paul). Baltimore; London: Johns Hopkins University Press.
ISBN 978-0-8018-6763-7. LCCN 2001000242 (https://lccn.loc.gov/2001000242). OCLC 1088130487 (htt
ps://www.worldcat.org/oclc/1088130487)..
Stewart, Tabori & Chang (1997). The Humongous Book of Dinosaurs. New York: Stewart, Tabori &
Chang. ISBN 978-1-55670-596-0. LCCN 97000398 (https://lccn.loc.gov/97000398). OCLC 1037269801
(https://www.worldcat.org/oclc/1037269801).
Sternberg, Charles Mortram (1966) [Original edition published by E. Cloutier, printer to the King, 1946].
Canadian Dinosaurs. Geological Series. Vol. 54 (2nd ed.). Ottawa: National Museum of Canada.
 LCCN gs46000214 (https://lccn.loc.gov/gs46000214). OCLC 1032865683 (https://www.worldcat.org/ocl
c/1032865683).

Notes
1. Dinosaurs (including birds) are members of the natural group Reptilia. Their biology does not precisely
correspond to the antiquated class Reptilia of Linnaean taxonomy, consisting of cold-blooded amniotes
without fur or feathers. As Linnean taxonomy was formulated for modern animals prior to the study of
evolution and paleontology, it fails to account for extinct animals with intermediate traits between
traditional classes.

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