Dinosaurs are a diverse group of reptiles[note

1] of the clade Dinosauria. They first appeared during the Triassic period, between 245 and
233.23 million years ago (mya), although the exact origin and timing of the evolution of
dinosaurs is a subject of active research. They became the 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 are all dinosaurs other than birds.

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 (birds)
and fossil remains. Through the first half of the 20th century, before birds were recognized as
dinosaurs, most of the scientific community believed dinosaurs to have been sluggish and cold-
blooded. Most research conducted 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 quadrupedal species,
and some were able to shift between these stances. 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 feet) and heights of 18 m (59 ft) and were the largest land
animals of all time. 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.

The first dinosaur fossils were recognized in the early 19th century, with the name "dinosaur"
(meaning "terrible lizard") being coined by Sir Richard Owen in 1841 to refer to these "great
fossil lizards". 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 and films, such as Jurassic Park. Persistent public
enthusiasm for the animals has resulted in significant funding for dinosaur science, and new
discoveries are regularly covered by the media.

Definition
Under phylogenetic nomenclature, dinosaurs are usually defined as the group consisting of the
most recent common ancestor (MRCA) of Triceratops and modern birds (Neornithes), and all its
descendants.[7] It has also been suggested that Dinosauria be defined with respect to the MRCA
of Megalosaurus and Iguanodon, because these were two of the three genera cited by Richard
Owen when he recognized the Dinosauria.[8] Both definitions result in the same set of animals
being defined as dinosaurs: "Dinosauria = Ornithischia + Saurischia". This definition includes
major groups such as ankylosaurians (armored herbivorous quadrupeds), stegosaurians (plated
herbivorous quadrupeds), ceratopsians (bipedal or quadrupedal herbivores with neck frills),
pachycephalosaurians (bipedal herbivores with thick skulls), ornithopods (bipedal or
quadrupedal herbivores including "duck-bills"), theropods (mostly bipedal carnivores and birds),
and sauropodomorphs (mostly large herbivorous quadrupeds with long necks and tails).[9]

Birds are now recognized as being the sole surviving lineage of theropod dinosaurs. In traditional
taxonomy, birds were considered a separate class that had evolved from dinosaurs, a distinct
superorder. However, a majority of contemporary paleontologists concerned with dinosaurs
reject the traditional style of classification in favor of phylogenetic taxonomy; this approach
requires that, for a group to be natural, all descendants of members of the group must be included
in the group as well. Birds are thus considered to be dinosaurs and dinosaurs are, therefore, not
extinct.[10] Birds are classified as belonging to the subgroup Maniraptora, which are
coelurosaurs, which are theropods, which are saurischians, which are dinosaurs.[11]

Research by Matthew G. Baron, David B. Norman, and Paul M. Barrett in 2017 suggested a
radical revision of dinosaurian systematics. Phylogenetic analysis by Baron et al. recovered the
Ornithischia as being closer to the Theropoda than the Sauropodomorpha, as opposed to the
traditional union of theropods with sauropodomorphs. They resurrected the clade Ornithoscelida
to refer to the group containing Ornithischia and Theropoda. Dinosauria itself was re-defined as
the last common ancestor of Triceratops horridus, Passer domesticus and Diplodocus carnegii,
and all of its descendants, to ensure that sauropods and kin remain included as dinosaurs.[12][13]

General description

Triceratops skeleton, Natural History Museum of Los Angeles County

Using one of the above definitions, dinosaurs can be generally described as archosaurs with hind
limbs held erect beneath the body.[14] Other prehistoric animals, including pterosaurs,
mosasaurs, ichthyosaurs, plesiosaurs, and Dimetrodon, while often popularly conceived of as
dinosaurs, are not taxonomically classified as dinosaurs.[15] Pterosaurs are distantly related to
dinosaurs, being members of the clade Ornithodira. The other groups mentioned are, like
dinosaurs and pterosaurs, members of Sauropsida (the reptile and bird clade), except Dimetrodon
(which is a synapsid). None of them had the erect hind limb posture characteristic of true
dinosaurs.[16]

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.[17] 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.[18][19][20] A 1995
study predicted that about 3,400 dinosaur genera ever existed, including many that would not
have been preserved in the fossil record.[21]

In 2016, the estimated number of dinosaur species that existed in the Mesozoic was 1,543–2,468.
[22][23] In 2021, the number of modern-day birds (avian dinosaurs) was estimated to be at
10,806 species.[24] 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.[25] 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.
[26]

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.[27]

Labeled diagram of a typical archosaur skull, the skull of Dromaeosaurus

A detailed assessment of archosaur interrelations by Sterling Nesbitt[28] 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 axis, the first two neck vertebrae
Apex of a deltopectoral crest (a projection on which the deltopectoral 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.[7]

Hip joints and hindlimb postures of: (left to right) typical reptiles (sprawling), dinosaurs and
mammals (erect), and rauisuchians (pillar-erect)
A variety of other skeletal features are shared by dinosaurs. However, because they are either
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.[29]
Additionally, several characteristics once thought to be synapomorphies are now known to have
appeared before dinosaurs, or were absent in the earliest dinosaurs and independently evolved by
different dinosaur groups. These include an elongated scapula, or shoulder blade; a sacrum
composed of three or more fused vertebrae (three are found in some other archosaurs, but only
two are found in Herrerasaurus);[7] and a perforate acetabulum, or hip socket, with a hole at the
center of its inside surface (closed in Saturnalia tupiniquim, for example).[30][31] 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.[32]

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.[33] 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.[34] Their erect posture enabled early
dinosaurs to breathe easily while moving, which likely permitted stamina and activity levels that
surpassed those of "sprawling" reptiles.[35] Erect limbs probably also helped support the
evolution of large size by reducing bending stresses on limbs.[36] 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.[36]

History of study
Further information: History of paleontology
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.
[37] Villagers in central China have long unearthed fossilized "dragon bones" for use in
traditional medicines.[38] In Europe, dinosaur fossils were generally believed to be the remains
of giants and other biblical creatures.[39]

Early dinosaur research

William Buckland
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,[40] 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).[41] 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.[42][43] 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 impicatum",[44][45] that had been found in Caswell, near Witney, Oxfordshire.[46]

Sir Richard Owen's coining of the word dinosaur, at a meeting of the British Association for the
Advancement of Science in 1841
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.[40][47] 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.[48][49]

The study of these "great fossil lizards" soon became of great interest to European and American
scientists, and in 1841 the English paleontologist Sir Richard Owen coined the term "dinosaur",
using it to refer to the "distinct tribe or sub-order of Saurian Reptiles" that were then being
recognized in England and around the world.[50][51] The term is derived from Ancient Greek
δεινός (deinos) 'terrible, potent or fearfully great', and σαῦρος (sauros) 'lizard or reptile'.[50][52]
Though the taxonomic name has often been interpreted as a reference to dinosaurs' teeth, claws,
and other fearsome characteristics, Owen intended it to also evoke their size and majesty.[53]
Owen recognized that the remains that had been found so far, Iguanodon, Megalosaurus and
Hylaeosaurus, shared a number of distinctive features, and so decided to present them as a
distinct taxonomic group. 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.[54]

Discoveries in North America

Edward Drinker Cope

Othniel Charles Marsh

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 discoveries sparked a wave of interests in dinosaurs in
the United States, known as dinosaur mania.[55]

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. Unfortunately,
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.[56]

"Dinosaur renaissance" and beyond

Main article: Dinosaur renaissance

John Ostrom's original restoration of Deinonychus, published in 1969

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.[57][58] At the end of the 1960s, however, the field of dinosaur research
experienced a surge in activity that remains ongoing.[59] Several seminal studies led to this
activity. First, John Ostrom discovered the bird-like dromaeosaurid theropod Deinonychus and
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.[57] 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),[60][61] which were
subsequently summarized in his 1986 book The Dinosaur Heresies.[62]

Paleontologist Robert T. Bakker with a mounted skeleton of a tyrannosaurid (Gorgosaurus

libratus)
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.[18] As of 2008, over 30 new species of dinosaurs were named each
year.[63] 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 named each year between
2009 and 2020. As a consequence, more sauropodomorphs were named between 1990 and 2020
than in all previous years combined.[64] 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.[63] Asian localities have produced the most complete
theropod specimens,[65] while North American localities have produced the most complete
sauropodomorph specimens.[64]

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.[66][67] Reference books summarizing the state of
dinosaur research, such as David B. Weishampel and colleagues' The Dinosauria, made
knowledge more accessible[68] 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.[69]

Soft tissue and molecular preservation

An Edmontosaurus specimen's skin impressions found in 1999

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.[70] 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.[71] 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.[72] Owen and Hooley subsequently described skin
impressions of Hypsilophodon and Iguanodon in 1885 and 1917.[71] Since then, scale
impressions have been most frequently found among hadrosaurids, where the impressions are
known from nearly the entire body across multiple specimens.[73]

Colour restoration of Sinosauropteryx

Colour restoration of Psittacosaurus

Starting from the 1990s, major discoveries of exceptionally preserved fossils in deposits known
as conservation Lagerstätten contributed to research on dinosaur soft tissues.[74][75] Chiefly
among these were the 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.[76][77] In living reptiles and mammals,
pigment-storing cellular structures known as melanosomes are partially responsible for
producing colouration.[78][79] 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.[80] This has enabled multiple full-body
reconstructions of dinosaur colouration, such as for Sinosauropteryx[81] and Psittacosaurus[82]
by Jakob Vinther and colleagues, and similar techniques have also been extended to dinosaur
fossils from other localities.[78] (However, some researchers have also suggested that fossilized
melanosomes represent bacterial remains.[83][84]) 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).[85][86] 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.[87]

Scipionyx fossil with intestines, Natural History Museum of Milan

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.[88][89][90]
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.[91] This discovery suggested that original
soft tissues could be preserved over geological time.[70] Later, in 2009, Schweitzer 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.[92] Both specimens yielded collagen protein
sequences that were viable for molecular phylogenetic analyses, which grouped them with birds
as would be expected.[92][93] The extraction of fragmentary DNA has also been reported for
both of these fossils,[94] along with a specimen of Hypacrosaurus.[95] 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.[96]
Suggestions that these structures represent bacterial biofilms[97] have been rejected,[98] but
cross-contamination remains a possibility that is difficult to detect.[99]

Evolutionary history
Origins and early evolution
Full skeleton of an early carnivorous dinosaur, displayed in a glass case in a museum
The early dinosaurs Herrerasaurus (large), Eoraptor (small) and a Plateosaurus skull, from the
Triassic
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.[100][101] The oldest dinosaur fossils known from substantial remains date to
the Carnian epoch of the Triassic period, which have been primarily found in the Ischigualasto
and Santa Maria Formations of Argentina, and the Pebbly Arkose Formation of Zimbabwe.[102]

The Ischigualasto Formation (radiometrically dated to be approximately 231-230 million years

old[103]) has produced the early saurischian Eoraptor, originally considered a member of the
Herrerasauridae[104] but now considered to be an early sauropodomorph, along with the
herrerasaurids Herrerasaurus and Sanjuansaurus, and the sauropodomorphs Chromogisaurus,
Eodromaeus, and Panphagia.[105] Eoraptor's likely resemblance to the common ancestor of all
dinosaurs suggests that the first dinosaurs would have been small, bipedal predators.[106][107]
[108] The Santa Maria Formation (radiometrically dated to be older, at 233.23 million years
old[109]) has produced the herrerasaurids Gnathovorax and Staurikosaurus, along with the
sauropodomorphs Bagualosaurus, Buriolestes, Guaibasaurus, Macrocollum, Nhandumirim,
Pampadromaeus, Saturnalia, and Unaysaurus.[105] 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.[102]

Less well-preserved remains of the sauropodomorphs Jaklapallisaurus and Nambalia, along with
the early saurischian Alwalkeria, are respectively known fron the Upper Maleri and Lower
Maleri Formations of India.[110] Meanwhile, 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.[105][111]
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 tell if it was a dinosaur or only a close
relative.[112] 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[113] to 21 million years.[109]

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.[114] 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,[115][116] and the exact date of their extinction is uncertain. These losses left
behind a land fauna of crocodylomorphs, dinosaurs, mammals, pterosaurians, and turtles.[7] 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.[9] Also notably,
there was a heightened rate of extinction during the Carnian pluvial event.[117]

Evolution and paleobiogeography

The supercontinent Pangaea in the early Mesozoic (around 200 million years ago)
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.[118] 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 so must have employed
other means of breaking down food farther along the digestive tract.[119] 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.[118] 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.[119] Another notable evolutionary event of the Jurassic was the appearance of true birds,
descended from maniraptoran coelurosaurians.[11]

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.[118] 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.[119] Some sauropods also evolved tooth batteries,
best exemplified by the rebbachisaurid Nigersaurus.[120]

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.[118] Flowering plants were greatly radiating,[119] with the first
grasses appearing by the end of the Cretaceous.[121] 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.
[119]

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.[122]

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.[123] Terror birds and bathornithids occupied carnivorous guilds alongside
predatory mammals,[124][125] and ratites are still fairly successful as mid-sized herbivores;
eogruiids similarly lasted from the Eocene to Pliocene, only becoming extinct very recently after
over 20 million years of co-existence with many mammal groups.[126]

Classification
Main article: Dinosaur classification

Saurischian pelvis structure (left side)

Tyrannosaurus pelvis (showing saurischian structure – left side)

Ornithischian pelvis structure (left side)

Edmontosaurus pelvis (showing ornithischian structure – left side)

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.[27]

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 pubis bone directed cranially, or forward.[34]
This basic form was modified by rotating the pubis backward to varying degrees in several
groups (Herrerasaurus,[127] therizinosauroids,[128] dromaeosaurids,[129] and birds[11]).
Saurischia includes the theropods (exclusively bipedal and with a wide variety of diets) and
sauropodomorphs (long-necked herbivores which include advanced, quadrupedal groups).[26]
[130]

By contrast, ornithischians—"bird-hipped", from the Greek ornitheios (ὀρνίθειος) meaning "of a

bird" and ischion (ἰσχίον) meaning "hip joint"—had a pelvis that superficially 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".[27]

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.[131] Further details and other hypotheses of
classification may be found on individual articles.

Dinosauria

Restoration of six ornithopods; far left: Camptosaurus, left: Iguanodon, center background:
Shantungosaurus, center foreground: Dryosaurus, right: Corythosaurus, far right (large)
Tenontosaurus.
†Ornithischia ("bird-hipped"; diverse bipedal and quadrupedal herbivores)
†Heterodontosauridae (small herbivores/omnivores with prominent canine-like teeth)
†Genasauria ("cheeked lizards")
†Thyreophora (armored dinosaurs; bipeds and quadrupeds)
†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)
Restoration of four ceratopsids: top left - Triceratops, top right - Styracosaurus, bottom left -
Anchiceratops, bottom right - Chasmosaurus.
†Pachycephalosauria (bipeds with domed or knobby growth on skulls)
†Ceratopsia (bipeds and quadrupeds; many had neck frills and horns)
†Chaoyangsauridae (small, frill-less basal ceratopsians)
†Neoceratopsia ("new ceratopsians")
†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)
†Triceratopsini (very large chasmosaurines with long 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)
†Camptosauridae (mid-sized, stocky)
†Styracosterna ("spiked sterna")
†Hadrosauriformes (ancestrally had a thumb spike)
†Hadrosauroidea (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)

Restoration of four macronarian sauropods: from left to right Camarasaurus, Brachiosaurus,

Giraffatitan, and Euhelopus
†Sauropodomorpha (herbivores with small heads, long necks, and long tails)
†Unaysauridae (primitive, strictly bipedal "prosauropods")
†Plateosauria (diverse; bipeds and quadrupeds)
†Massospondylidae (long-necked, primitive sauropodomorphs)
†Riojasauridae (large, primitive sauropodomorphs)
†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.[87][91] 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).[132][133] Two topics of particular interest and study
have been dinosaur size and behavior.[134]

Size
Main article: Dinosaur size

Scale diagram comparing the average human to the longest known dinosaurs in five major
clades:
Sauropoda (Supersaurus vivianae)
Ornithopoda (Shantungosaurus giganteus)
Theropoda (Spinosaurus aegyptiacus)
Thyreophora (Stegosaurus ungulatus)
Marginocephalia (Triceratops prorsus)
Current evidence suggests that dinosaur average size varied through the Triassic, Early Jurassic,
Late Jurassic and Cretaceous.[107] Predatory theropod dinosaurs, which occupied most
terrestrial carnivore niches during the Mesozoic, most often fall into the 100 to 1000 kg (220 to
2200 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.
[135] The mode of Mesozoic dinosaur body masses is between 1 to 10 metric tons (1.1 to 11.0
short tons).[136] 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).[137]

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.[138] 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 them to subsist on food with lower
nutritive value than smaller animals. Sauropod remains are mostly found in rock formations
interpreted as dry or seasonally dry, and the ability to eat large quantities of low-nutrient browse
would have been advantageous in such environments.[139]

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.[140]

Comparative size of Argentinosaurus to the average human

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;[141] this mount is 12 meters
(39 ft) tall and 21.8 to 22.5 meters (72 to 74 ft) long,[142][143] and would have belonged to an
animal that weighed between 30000 and 60000 kilograms (70000 and 130000 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.
[144] 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.[145]

An adult bee hummingbird, the smallest known dinosaur

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 80000 to
100000 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[139] (formerly
Seismosaurus), the 33-to-34-meter (108 to 112 ft) long Supersaurus,[146] 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 122400 kg (270000 lb).[139] However, as no further evidence of sauropods of
this size has been found, and the discoverer, Cope, had made typographic errors before, it is
likely to have been an extreme overestimation.[147]

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).[148][149] Other large
carnivorous theropods included Giganotosaurus, Carcharodontosaurus and Tyrannosaurus.[149]
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).[150] The largest individuals may have weighed as much as 16 metric tons
(18 short tons).[151]

The smallest dinosaur known is the bee hummingbird,[152] with a length of only 5 centimeters
(2.0 in) and mass of around 1.8 g (0.063 oz).[153] The smallest known non-avialan dinosaurs
were about the size of pigeons and were those theropods most closely related to birds.[154] 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)[155] and a total skeletal length of 34
centimeters (1.12 ft).[154][155] The smallest herbivorous non-avialan dinosaurs included
Microceratus and Wannanosaurus, at about 60 centimeters (2.0 ft) long each.[156][157]

Behavior

A nesting ground of the hadrosaur Maiasaura peeblesorum was discovered in 1978

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.[132]
The first potential evidence for herding or flocking as a widespread behavior 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 Bernissart, Belgium, after they fell into a deep,
flooded sinkhole and drowned.[158] 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,[159] although there is no evidence for
specific herd structures.[160] 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.[161] The interpretation
of dinosaurs as gregarious has also extended to depicting carnivorous theropods as pack hunters
working together to bring down large prey.[162][163] 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.[164]

Restoration of two Centrosaurus apertus engaged in intra-specific combat

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.[165]

From a behavioral standpoint, one of the most valuable dinosaur fossils was discovered in the
Gobi Desert in 1971. It included a Velociraptor attacking a Protoceratops,[166] providing
evidence that dinosaurs did indeed attack each other.[167] 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.[167] Cannibalism
amongst some species of dinosaurs was confirmed by tooth marks found in Madagascar in 2003,
involving the theropod Majungasaurus.[168]
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.[169]

Based on fossil evidence from dinosaurs such as Oryctodromeus, some ornithischian species
seem to have led a partially fossorial (burrowing) lifestyle.[170] Many modern birds are arboreal
(tree climbing), and this was also true of many Mesozoic birds, especially the enantiornithines.
[171] While some early bird-like species may have already been arboreal as well (including
dromaeosaurids) such as Microraptor[172]) 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,[132] whether diplodocids could create sonic booms via whip-like tail
snapping,[173] and whether sauropods could float.[174]

Communication

Restoration of a striking and unusual visual display in a Lambeosaurus magnicristatus

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.[175] Reconstruction of the plumage color of Anchiornis, suggest the importance of
color in visual communication in non-avian dinosaurs.[176]

The evolution of dinosaur vocalization is less certain. The earliest remains of the avian 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.[177] Paleontologist Phil Senter has
suggested that non-avian dinosaurs relied 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). He states they were unlikely to have been capable of vocalizing
since their closest relatives, crocodilians and birds, use different means to vocalize, the larynx
and syrinx respectively, suggesting they evolved independently and their common ancestor was
mute.[175] Other researchers have countered that vocalizations also exist in turtles, the closest
relatives of archosaurs, suggesting that the trait is ancestral to their lineage.[178]

In 2023, a fossilized larynx was described from a specimen of the ankylosaurid Pinacosaurus.
The structure was composed of cricoid and arytenoid cartilages, similar to those of non-avian
reptiles. However, the mobile cricoid-arytenoid joint and long arytenoid cartilages would have
allowed for air-flow control similar to 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.[179] 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.[180] The crests of the Lambeosaurini and nasal chambers of ankylosaurids have been
suggested to have functioned in acoustic resonance.[181][182]

Reproductive biology
See also: Dinosaur egg
Three bluish eggs with black speckling sit atop a layer of white mollusk shell pieces, surrounded
by sandy ground and small bits of bluish stone
Nest of a plover (Charadrius)
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.
[183] 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.[184]

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 medullary tissue is a general characteristic of all
dinosaurs.[185]

Fossil interpreted as a nesting oviraptorid Citipati at the American Museum of Natural History.
Smaller fossil far right showing inside one of the eggs.
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.[186] A specimen of the oviraptorid Citipati
osmolskae was discovered in a chicken-like brooding position in 1993,[187] which may indicate
that they had begun using an insulating layer of feathers to keep the eggs warm.[188] An embryo
of the basal sauropodomorph Massospondylus was found without teeth, indicating that some
parental care was required to feed the young dinosaurs.[189] Trackways have also confirmed
parental behavior among ornithopods from the Isle of Skye in northwestern Scotland.[190]

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.[191][192][193][194] Both Tyrannosaurus and Troodon had juveniles with
clear superprecociality and likely occupying different ecological niches than the adults.[184]
Superprecociality has been inferred for sauropods.[195]

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.[196]

Physiology
Main article: Physiology of dinosaurs
Because both modern crocodilians and birds have four-chambered hearts (albeit