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

They first appeared

during the Triassic period, between 243 and 233.23 million years ago, [1][2] although the exact
origin and timing of the evolution of dinosaurs is the subject of active research. [3] They became
the dominant terrestrial vertebrates after the Triassic–Jurassic extinction event 201 million years
ago; their dominance continued through the Jurassic and Cretaceous periods. Reverse genetic
engineering[4] and the fossil record both demonstrate that birds are modern feathered
dinosaurs,[5] having evolved from earlier theropods during the late Jurassic Period.[6] As such,
birds were the only dinosaur lineage to survive the Cretaceous–Paleogene extinction event 66
million years ago.[7] Dinosaurs can therefore be divided into avian dinosaurs, or birds; and non-
avian dinosaurs, which are all dinosaurs other than birds. This article deals primarily with non-
avian dinosaurs.
Dinosaurs are a varied group of animals
from taxonomic, morphological and ecological standpoints. Birds, at over 10,000 living
species,[8]are the most diverse group of vertebrates besides perciform fish.[9] Using fossil
evidence, paleontologists have identified over 500 distinct genera[10] and more than 1,000
different species of non-avian dinosaurs.[11] Dinosaurs are represented on every continent by
both extantspecies (birds) and fossil remains.[12] Through the first half of the 20th century, before
birds were recognized to be 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 all dinosaurs were active animals with elevated metabolisms and numerous
adaptations for social interaction. Some were herbivorous, others carnivorous. Evidence
suggests that egg-laying and nest-building are additional traits shared by all dinosaurs, avian and
non-avian alike.
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) [13] and heights of 18 meters (59 feet)[14] and were the
largest land animals of all time. Still, the idea that non-avian dinosaurs were uniformly gigantic is
a misconception based in part on preservation bias, as large, sturdy bones are more likely to last
until they are fossilized. Many dinosaurs were quite small: Xixianykus, for example, was only
about 50 cm (20 in) long.
Since the first dinosaur fossils were recognized in the early 19th century, mounted fossil dinosaur
skeletons have been major attractions at museums around the world, and dinosaurs have
become an enduring part of world culture. The large sizes of some dinosaur groups, as well as
their seemingly monstrous and fantastic nature, have ensured dinosaurs' 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.

Contents

 1Etymology
 2Definition
o 2.1General description
o 2.2Distinguishing anatomical features
 3Evolutionary history
o 3.1Origins and early evolution
o 3.2Evolution and paleobiogeography
 4Classification
o 4.1Taxonomy
 5Biology
o 5.1Size
 5.1.1Largest and smallest
o 5.2Behavior
o 5.3Communication
o 5.4Reproductive biology
o 5.5Physiology
 6Origin of birds
o 6.1Feathers
o 6.2Skeleton
o 6.3Soft anatomy
o 6.4Behavioral evidence
 7Extinction of major groups
o 7.1Impact event
o 7.2Deccan Traps
o 7.3Possible Paleocene survivors
 8History of study
o 8.1"Dinosaur renaissance"
o 8.2Soft tissue and DNA
 9Cultural depictions
 10See also
 11Notes
 12References
 13Further reading
 14External links

Etymology
The taxon 'Dinosauria' was formally named in 1841 by paleontologist Sir Richard Owen, who
used it to refer to the "distinct tribe or sub-order of Saurian Reptiles" that were then being
recognized in England and around the world.[15] The term is derived from Ancient
Greek δεινός (deinos), meaning 'terrible, potent or fearfully great', and σαῦρος (sauros), meaning
'lizard or reptile'.[15][16] Though the taxonomic name has often been interpreted as a reference to
dinosaurs' teeth, claws, and other fearsome characteristics, Owen intended it merely to evoke
their size and majesty.[17]
Other prehistoric animals, including pterosaurs, mosasaurs, ichthyosaurs, plesiosaurs,
and Dimetrodon, while often popularly conceived of as dinosaurs, are
not taxonomically classified as dinosaurs.[18] 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).

Definition
Triceratops skeleton, Natural History Museum of Los Angeles County

Under phylogenetic nomenclature, dinosaurs are usually defined as the group consisting of
the most recent common ancestor (MRCA) of Triceratops and Neornithes, and all its
descendants.[19] 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.[20] Both definitions result in the same set of animals
being defined as dinosaurs: "Dinosauria = Ornithischia + Saurischia",
encompassing ankylosaurians(armored herbivorous quadrupeds), stegosaurians (plated
herbivorous quadrupeds), ceratopsians (herbivorous quadrupeds with horns and
frills), 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).[21]
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.[22] Birds are classified as belonging to the subgroup Maniraptora, which
are coelurosaurs, which are theropods, which are saurischians, which are dinosaurs.[23]
Research by Matthew 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 Ornithoscelidato refer to the group containing Ornithischia and Theropoda. Dinosauria itself
was re-defined as the last common ancestor of Triceratops horridus, Passer
domesticus, Diplodocus carnegii, and all of its descendants, to ensure that sauropods and kin
remain included as dinosaurs.[24][25]

General description
In phylogenetic taxonomy, birds are included in the group Dinosauria.

Using one of the above definitions, dinosaurs can be generally described as archosaurs with hind
limbs held erect beneath the body.[26] Many prehistoric animal groups are popularly conceived of
as dinosaurs, such as ichthyosaurs, mosasaurs, plesiosaurs, pterosaurs,
and pelycosaurs(especially Dimetrodon), but are not classified scientifically as dinosaurs, and
none had the erect hind limb posture characteristic of true dinosaurs.[27]Dinosaurs were the
dominant terrestrial vertebrates of the Mesozoic, 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. [28]
Dinosaurs have always been an extremely varied group of animals; according to a 2006 study,
over 500 non-avian dinosaur genera have been identified with certainty so far, 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.[10] An earlier study predicted that about 3,400 dinosaur genera
existed, including many that would not have been preserved in the fossil record. [29] By September
17, 2008, 1,047 different species of dinosaurs had been named.[11]
In 2016, the estimated number of dinosaur species that existed in the Mesozoic era was
estimated to be 1,543–2,468.[30][31] 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
period.[12] 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.[32]

Distinguishing anatomical features

While recent discoveries have made it more difficult to present a universally agreed-upon list of
dinosaurs' distinguishing features, nearly all dinosaurs discovered so far share certain
modifications to the ancestral archosaurian skeleton, or are clear 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.[33]
A detailed assessment of archosaur interrelations by Sterling Nesbitt[34] 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, present in the
anterior (front) neck vertebrae behind the atlas and axis, the first two neck vertebrae
 apex of 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 (thighbone) 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.[19]

Diagram of a typical diapsid skull

j: jugal bone, po: postorbital bone, p: parietal bone, sq: squamosal bone, q: quadrate bone,
qj: quadratojugal bone

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 temporal fenestrae (openings in the skull behind the eyes), and as members of the
diapsid group Archosauria, had additional openings in the snoutand lower jaw.[35] 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);[19] and a perforate acetabulum, or hip socket, with a hole at the center of its
inside surface (closed in Saturnalia, for example).[36][37] Another difficulty of determining distinctly
dinosaurian features is that early dinosaurs and other archosaurs from the late Triassic are often
poorly known and were similar in many ways; these animals have sometimes been misidentified
in the literature.[38]

Hip joints and hindlimb postures of: (left to right) typical reptiles (sprawling), dinosaurs and mammals
(erect), and rauisuchians (erect)

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.[39] 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.[40] Their erect posture enabled early
dinosaurs to breathe easily while moving, which likely permitted stamina and activity levels
that surpassed those of "sprawling" reptiles.[41] Erect limbs probably also helped support the
evolution of large size by reducing bending stresses on limbs.[42] 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.[42]

Evolutionary history
Main article: Evolution of dinosaurs

Origins and early evolution

Dinosaurs diverged from their archosaur ancestors during the middle to late Triassic period,
roughly 20 million years after the Permian–Triassic extinction event wiped out an estimated 95%
of all life on Earth.[43][44] Radiometric dating of the rock formation that contained fossils from the
early dinosaur genus Eoraptor at 231.4 million years old establishes its presence in the fossil
record at this time.[45] Paleontologists think that Eoraptor resembles the common ancestor of all
dinosaurs;[46] if this is true, its traits suggest that the first dinosaurs were small,
bipedal predators.[47] The discovery of primitive, dinosaur-like ornithodirans such
as Marasuchus and Lagerpeton in Argentinian Middle Triassic strata supports this view; analysis
of recovered fossils suggests that these animals were indeed small, bipedal predators. Dinosaurs
may have appeared as early as 243 million years ago, as evidenced by remains of the
genus Nyasasaurus from that period, though known fossils of these animals are too fragmentary
to tell if they are dinosaurs or very close dinosaurian relatives.[48] Recently, it has been
determined that Staurikosaurusfrom the Santa Maria Formation dates to 233.23 Ma, making it
older in geologic age than Eoraptor.[1]
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 pseudosuchia, such
as aetosaurs, ornithosuchids and rauisuchians, which were more successful than the
dinosaurs.[49] 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 200 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, respectively,[50][51] and the exact date of their extinction is uncertain.
These losses left behind a land fauna of crocodylomorphs, dinosaurs, mammals, pterosaurians,
and turtles.[19] 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.[21] Also notably, there was a heightened rate of extinction during the Carnian
Pluvial Event.[52]

Evolution and paleobiogeography

Dinosaur evolution after the Triassic follows 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.[53] 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.[54] The general homogeneity of dinosaurian faunas continued into the middle and late
Jurassic, where most localities had predators consisting of ceratosaurians, spinosauroids,
and carnosaurians, and herbivores consisting of stegosaurian ornithischians and large
sauropods. Examples of this include the Morrison Formationof North America and Tendaguru
Beds of Tanzania. Dinosaurs in China show some differences, with
specialized sinraptorid theropods and unusual, long-necked sauropods
like Mamenchisaurus.[53] Ankylosaurians and ornithopods were also becoming more common,
but prosauropods had become extinct. Conifers and pteridophytes were the most common
plants. Sauropods, like the earlier prosauropods, 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.[54] Another notable evolutionary
event of the Jurassic was the appearance of true birds, descended
from maniraptorancoelurosaurians.[55]

Skeleton of Marasuchus lilloensis, a dinosaur-like ornithodiran

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.[53] 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 tooth batteries,
taken to its extreme in hadrosaurids.[54] Some sauropods also evolved tooth batteries, best
exemplified by the rebbachisaurid Nigersaurus.[56]

The early forms Herrerasaurus(large), Eoraptor (small) and a Plateosaurus skull

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 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.[53] Flowering plants were greatly radiating,[54] with the first grasses
appearing by the end of the Cretaceous.[57] Grinding hadrosaurids and shearing ceratopsians
became extremely diverse across North America and Asia. Theropods were also radiating as
herbivores or omnivores, with therizinosaurians and ornithomimosaurians becoming common.[54]
The Cretaceous–Paleogene extinction event, which occurred approximately 66 million years ago
at the end of the Cretaceous period, caused the extinction of all dinosaur groups except for
the neornithine birds. Some other diapsid groups, such as
crocodilians, sebecosuchians, turtles, lizards, snakes, sphenodontians, and choristoderans, also
survived the event.[58]
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 Paleogeneperiod,
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 cited 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.[59] Terror birds and bathornithids occupied carnivorous guilds alongside predatory
mammals,[60][61] 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.[62]

Classification
Main article: Dinosaur 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.[33]
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.[40] This basic form was modified by rotating the pubis backward to varying degrees in
several groups (Herrerasaurus,[63]therizinosauroids,[64] dromaeosaurids,[65] and birds[55]). Saurischia
includes the theropods (exclusively bipedal and with a wide variety of diets)
and sauropodomorphs (long-necked herbivoreswhich include advanced, quadrupedal
groups).[66][67]
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
which were primarily herbivores. (NB: the terms "lizard hip" and "bird hip" are misnomers – birds
evolved from dinosaurs with "lizard hips".)[33]

Saurischian pelvis structure (left side)

 

Tyrannosaurus pelvis (showing saurischian structure – left side)

Ornithischian pelvis structure (left side)

Edmontosaurus pelvis (showing ornithischian structure – left side)

Taxonomy
The following is a simplified classification of dinosaur groups based on their evolutionary
relationships, and organized based on the list of Mesozoic dinosaur species provided by Holtz
(2007).[7] A more detailed version can be found at Dinosaur classification. The dagger (†) is used
to signify groups with no living members.

 Dinosauria

 Saurischia ("lizard-hipped"; includes Theropoda and Sauropodomorpha)

 †Herrerasauria (early bipedal carnivores)

 Theropoda (all bipedal; most were carnivorous)
Artist's impression of six dromaeosaurid theropods: from left to
right Microraptor, Velociraptor, Austroraptor, Dromaeosaurus, Utahraptor, and Deinonychus

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

 †Dilophosauridae (early crested and carnivorous theropods)
 †Ceratosauria (generally elaborately horned, the dominant southern carnivores of the
Cretaceous)
 Tetanurae ("stiff tails"; includes most theropods)

 †Megalosauroidea (early group of large carnivores including the semiaquatic

spinosaurids)
 †Carnosauria (Allosaurus and close relatives, like Carcharodontosaurus)
 Coelurosauria (feathered theropods, with a range of body sizes and niches) [5]

 †Compsognathidae (common early coelurosaurs with reduced forelimbs)

 †Tyrannosauridae (Tyrannosaurus and close relatives; had reduced forelimbs)
 †Ornithomimosauria ("ostrich-mimics"; mostly toothless; carnivores to possible
herbivores)
 †Alvarezsauroidea (small insectivores with reduced forelimbs each bearing one
enlarged claw)
 Maniraptora ("hand snatchers"; had long, slender arms and fingers)

 †Therizinosauria (bipedal herbivores with large hand claws and small heads)
 †Oviraptorosauria (mostly toothless; their diet and lifestyle are uncertain)
 †Archaeopterygidae (small, winged theropods or primitive birds)
 †Deinonychosauria (small- to medium-sized; bird-like, with a distinctive toe claw)
 Avialae (modern birds and extinct relatives)

 †Scansoriopterygidae (small primitive avialans with long third fingers)

 †Omnivoropterygidae (large, early short-tailed avialans)
 †Confuciusornithidae (small toothless avialans)
 †Enantiornithes (primitive tree-dwelling, flying avialans)
 Euornithes (advanced flying birds)

 †Yanornithiformes (toothed Cretaceous Chinese birds)

 †Hesperornithes (specialized aquatic diving birds)
 Aves (modern, beaked birds and their extinct relatives)

Artist's impression of four macronarian sauropods: from left to

right Camarasaurus, Brachiosaurus, Giraffatitan, and Euhelopus
 †Sauropodomorpha (herbivores with small heads, long necks, long tails)

 †Guaibasauridae (small, primitive, omnivorous sauropodomorphs)

 †Plateosauridae (primitive, strictly bipedal "prosauropods")
 †Riojasauridae (small, primitive sauropodomorphs)
 †Massospondylidae (small, primitive sauropodomorphs)
 †Sauropoda (very large and heavy, usually over 15 m (49 ft) long; quadrupedal)

 †Vulcanodontidae (primitive sauropods with pillar-like limbs)

 †Eusauropoda ("true sauropods")

 †Cetiosauridae ("whale reptiles")

 †Turiasauria (European group of Jurassic and Cretaceous sauropods)
 †Neosauropoda ("new sauropods")

 †Diplodocoidea (skulls and tails elongated; teeth typically narrow and pencil-like)
 †Macronaria (boxy skulls; spoon- or pencil-shaped teeth)

 †Brachiosauridae (long-necked, long-armed macronarians)

 †Titanosauria (diverse; stocky, with wide hips; most common in the late Cretaceous
of southern continents)

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 basal ornithopod herbivores/omnivores with

prominent canine-like teeth)
 †Thyreophora (armored dinosaurs; mostly quadrupeds)

 †Ankylosauria (scutes as primary armor; some had club-like tails)

 †Stegosauria (spikes and plates as primary armor)

 †Neornithischia ("new ornithischians")

 †Ornithopoda (various sizes; bipeds and quadrupeds; evolved a method of chewing

using skull flexibility and numerous teeth)
 †Marginocephalia (characterized by a cranial growth)

 †Pachycephalosauria (bipeds with domed or knobby growth on skulls)

 †Ceratopsia (quadrupeds with frills; many also had horns)

Biology
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 soft tissues.[68][69] 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).[70][71] Two topics of particular
interest and study have been dinosaur size and behavior.[72]

Size
Main article: Dinosaur size

Scale diagram comparing the average human to the largest known dinosaurs in five
major clades: Sauropoda (Argentinosaurus
huinculensis), Ornithopoda (Shantungosaurus giganteus), Theropoda(Spinosaurus
aegyptiacus), Thyreophora (Stegosaurus armatus) and Marginocephalia (Triceratops
prorsus)

Current evidence suggests that dinosaur average size varied through the
Triassic, early Jurassic, late Jurassic and Cretaceous periods.[46] 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.[73] The mode of Mesozoic dinosaur body masses is between
one and ten metric tonnes.[74] 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).[75]
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 were 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.[76] 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.[14]
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 ever
fossilize, 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.[77]

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 Berlin;[78] this mount is 12 meters (39 ft)
tall and 21.8–22.5 meters (72–74 ft) long,[79][80] 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
feet) long Diplodocus, which was discovered in Wyoming in the United
States and displayed in Pittsburgh's Carnegie Natural History Museum in
1907.[81] The longest dinosaur known from good fossil material is the Patagotitan:
the skeleton mount in the American Museum of Natural History is 37 meters
(121 ft) long. The Carmen Funes Museum has
an Argentinosaurus reconstructed skeleton mount 39.7 metres (130 ft) long.[82]

Comparative size of Eoraptor to the average human

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 length of 30–40 metres (98–131 ft); some of
the longest were the 33.5 meters (110 ft) long Diplodocus
hallorum[14] (formerly Seismosaurus), the 33–34 meters (108–112 ft)
long Supersaurus[83] and 37 metres (121 ft) long Patagotitan; and the tallest, the
18 meters (59 ft) tall Sauroposeidon, which could have reached a sixth-floor
window. The heaviest and longest dinosaur may have been Amphicoelias
fragillimus, 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).[14] However, as no
further evidence of sauropods of this size has been found, and the discoverer,
Edward Cope, had made typographic errors before, it is likely to have been an
extreme overestimation.[84]
The largest carnivorous dinosaur was Spinosaurus, reaching a length of 12.6 to
18 meters (41 to 59 ft), and weighing 7–20.9 tonnes (7.7–23 short
tons).[85][86] Other large carnivorous theropods
included Giganotosaurus, Carcharodontosaurus and Tyrannosaurus.[86] Therizin
osaurus and Deinocheirus were among the tallest of the theropods. The
largest Ornithischian dinosaur was probably
the hadrosaurid Shantungosaurus which measured 16 metres (52 ft) and
weighed about 13 tonnes (29,000 lb).
The smallest dinosaur known is the bee hummingbird,[87] with a length of only
5 cm (2.0 in) and mass of around 1.8 g (0.063 oz).[88] The smallest known non-
avialan dinosaurs were about the size of pigeons and were those theropods
most closely related to birds.[89] For example, Anchiornis huxleyi is currently the
smallest non-avialan dinosaur described from an adult specimen, with an
estimated weight of 110 grams[90] and a total skeletal length of 34 cm
(1.12 ft).[89][90] The smallest herbivorous non-avialan dinosaurs
included Microceratus and Wannanosaurus, at about 60 cm (2.0 ft) long
each.[7][91]

Behavior

A nesting ground of 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 (birds' closest living relatives), 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.[70]
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 bernissartensis, ornithischians that were then thought to have
perished together in Bernissart, Belgium, after they fell into a deep,
flooded sinkhole and drowned.[92] 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-bills(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,[93] although there is no
evidence for specific herd structures.[94] 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 the remains of over 20 Sinornithomimus, from one to
seven years old. This assemblage is interpreted as a social group that was
trapped in mud.[95] The interpretation of dinosaurs as gregarious has also
extended to depicting carnivorous theropods as pack hunters working together
to bring down large prey.[96][97] 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.[98]

Artist's rendering 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.[99]
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,[100] providing evidence that dinosaurs did indeed attack each
other.[101] 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.[101] Cannibalism amongst some species of dinosaurs was confirmed by
tooth marks found in Madagascar in 2003, involving the
theropod Majungasaurus.[102]
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.[103]
Based on current fossil evidence from dinosaurs such as Oryctodromeus, some
ornithischian species seem to have led a partially fossorial (burrowing)
lifestyle.[104] Many modern birds are arboreal (tree climbing), and this was also
true of many Mesozoic birds, especially the enantiornithines.[105] While some
early bird-like species may have already been arboreal as well
(including dromaeosaurids such as Microraptor[106]) 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,[107] whether diplodocids could create sonic booms via whip-
like tail snapping,[108] and whether sauropods could float.[109]

Communication
 Artist's impression 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.[110]Reconstruction
of the plumage color of Anchiornis huxleyi, suggest the importance of color in
visual communication in non-avian dinosaurs.[111] The evolution of dinosaur
vocalization is less certain. Paleontologist Phil Senter suggests 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 former via the larynx and the latter
through the unique syrinx, suggesting they evolved independently and their
common ancestor was mute.[110]
The earliest remains of a syrinx, which has enough mineral content for
fossilization, was found in a specimen of the duck-like Vegavis iaai dated 69-66
million year ago, and this organ is unlikely to have existed in non-avian
dinosaurs. However, in contrast to Senter, the researchers have suggested that
dinosaurs could vocalize and that the syrinx-based vocal system of birds
evolved from a larynx-based one, rather than the two systems evolving
independently.[112] A 2016 study suggests that dinosaurs produced closed mouth
vocalizations like cooing, which occur in both crocodilians and birds as well as
other reptiles. Such vocalizations evolved independently in extant archosaurs
numerous times, following increases in body size.[113] The crests of
the Lambeosaurini and nasal chambers of ankylosaurids have been suggested
to function in vocal resonance,[114][115] though Senter states that the presence of
resonance chambers in some dinosaurs is not necessarily evidence of
vocalization as modern snakes have such chambers which intensify their
hisses.[110]

Reproductive biology
See also: Dinosaur egg
Nest of a plover (Charadrius)

All dinosaurs lay amniotic eggs with hard shells made mostly of calcium
carbonate.[116] Eggs are usually laid in a nest. Most species create somewhat
elaborate nests, which can be cups, domes, plates, beds scrapes, mounds, or
burrows.[117] 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. [118]
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 rex 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.[119]

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.[120] A specimen of the Mongolian oviraptorid Citipati
osmolskae was discovered in a chicken-like brooding position in 1993,[121] which
may indicate that they had begun using an insulating layer of feathers to keep
the eggs warm.[122] A dinosaur embryo (pertaining to
the prosauropod Massospondylus) was found without teeth, indicating that some
parental care was required to feed the young dinosaurs.[123]Trackways have also
confirmed parental behavior among ornithopods from the Isle of Skye in
northwestern Scotland.[124]
However, there is ample evidence of supreprecociality 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 behaviour and the ability to fly soon after
birth.[125][126][127] Both Tyrannosaurus rex and Troodon formosus display juveniles
with clear supreprecociality and likely occupying different ecological niches than
the adults.[128] Superprecociality has been inferred for sauropods.[129]

Physiology
Main article: Physiology of dinosaurs
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.[130]While all modern birds have high metabolisms and are
"warm blooded" (endothermic), a vigorous debate has been ongoing since the
1960s regarding how far back in the dinosaur lineage this trait extends.
Scientists disagree as to whether non-avian dinosaurs were endothermic,
ectothermic, or some combination of both.[131]
After non-avian dinosaurs were discovered, paleontologists first posited that
they were ectothermic. This supposed "cold-bloodedness" 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 and sluggish remained a prevalent view until Robert T. "Bob"
Bakker, an early proponent of dinosaur endothermy, published an influential
paper on the topic in 1968.[132]
Modern evidence indicates that even non-avian dinosaurs and birds thrived in
cooler temperate climates, and that at least some early species must have
regulated their body temperature by internal biological means (aided by the
animals' bulk in large species and feathers or other body coverings in smaller
species). Evidence of endothermy in Mesozoic dinosaurs includes the discovery
of polar dinosaurs in Australia and Antarctica as well as analysis of blood-vessel
structures within fossil bones that are typical of endotherms. Scientific debate
continues regarding the specific ways in which dinosaur temperature regulation
evolved.[133][134]

Comparison between the air sacs of an abelisaur and a bird

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.[135] Early avian-style respiratory systems with air sacs may have
been capable of sustaining higher activity levels than those of mammals of
similar size and build. In addition to providing a very efficient supply of oxygen,
the rapid airflow would have been an effective cooling mechanism, which is
essential for animals that are active but too large to get rid of all the excess heat
through their skin.[136]
Like other reptiles, dinosaurs are primarily uricotelic, that is, their kidneys extract
nitrogenous wastes from their bloodstream and excrete it as uric acidinstead
of urea or ammonia via the ureters into the intestine. In most living species, uric
acid is excreted along with feces as a semisolid waste.[137][138][139] However, at
least some modern birds (such as hummingbirds) can be
facultatively ammonotelic, excreting most of the nitrogenous wastes as
ammonia.[140] They also excrete creatine, rather than creatinine like
mammals.[141] This material, as well as the output of the intestines, emerges from
the cloaca.[142][143] In addition, many species regurgitate pellets, and fossil pellets
that may have come from dinosaurs are known from as long ago as the
Cretaceous period.[144]

Origin of birds
Main article: Origin of birds
The possibility that dinosaurs were the ancestors of birds was first suggested in
1868 by Thomas Henry Huxley.[145] 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 their being descendants of generalized thecodonts, with
the key piece of evidence being the supposed lack of clavicles in
dinosaurs.[146] However, as later discoveries showed, clavicles (or a single
fused wishbone, which derived from separate clavicles) were not actually
absent;[55] they had been found as early as 1924 in Oviraptor, but misidentified
as an interclavicle.[147] In the 1970s, John Ostrom revived the dinosaur–bird
theory,[148] which gained momentum in the coming decades with the advent of
cladistic analysis,[149] and a great increase in the discovery of small theropods
and early birds.[35] Of particular note have been the fossils of the Yixian
Formation, where a variety of theropods and early birds have been found, often
with feathers of some type.[5][55] Birds share over a hundred distinct anatomical
features with theropod dinosaurs, which are now generally accepted to have
been their closest ancient relatives.[150] They are most closely allied
with maniraptoran coelurosaurs.[55] 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,[151] or that
maniraptoran theropods are the ancestors of birds but themselves are not
dinosaurs, only convergent with dinosaurs.[152]

Feathers
Main article: Feathered dinosaurs
 Various feathered non-avian dinosaurs,
including Archaeopteryx, Anchiornis, Microraptor and Zhenyuanlong

Feathers are one of the most recognizable characteristics of modern birds, and
a trait that was shared by all other dinosaur groups. 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.[153] Direct fossil evidence of feathers or feather-like structures has been
discovered in a diverse array of species in many non-avian dinosaur
groups,[5] both among saurischians and ornithischians. Simple, branched,
feather-like structures are known from heterodontosaurids,
primitive neornithischians[154] and theropods,[155] 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.[55][156] Feather-like
structures known as pycnofibres have also been found
in pterosaurs,[157] suggesting the possibility that feather-like filaments may have
been common in the bird lineage and evolved before the appearance of
dinosaurs themselves.[153] Research into the genetics of American alligators has
also revealed that crocodylian scutes do possess feather-keratins during
embryonic development, but these keratins are not expressed by the animals
before hatching.[158]
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 Darwin's
seminal The Origin of Species, 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 Compsognathus.[159] 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. Most of these specimens were unearthed in the lagerstätte of the Yixian
Formation, Liaoning, northeastern China, which was part of an island continent
during the Cretaceous. Though feathers have been found in only a few
locations, it is possible that non-avian dinosaurs elsewhere in the world were
also feathered. The lack of widespread fossil evidence for feathered non-avian
dinosaurs may be because delicate features like skin and feathers are not often
preserved by fossilization and thus are absent from the fossil record.[160]
The description of feathered dinosaurs has not been without controversy;
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,[161][162][163] and that maniraptoran dinosaurs with vaned feathers were not
actually dinosaurs, but convergent with dinosaurs.[152][162] 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.[164]
In 2016, it was reported that a dinosaur tail with feathers had been found
enclosed in amber. The fossil is about 99 million years old.[5][165][166]

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.[167]

Soft anatomy

Pneumatopores on the left ilium of Aerosteon riocoloradensis

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.[168] 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.[169][170]

Behavioral evidence
Fossils of the troodonts Mei and Sinornithoides demonstrate that some
dinosaurs slept with their heads tucked under their arms.[171] This behavior, which
may 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.[172] 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.[173]
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.[174]

Extinction of major groups

Main article: Cretaceous–Paleogene extinction event
The discovery that birds are a type of dinosaur showed that dinosaurs in general
are not, in fact, extinct as is commonly stated.[175] However, all non-avian
dinosaurs, estimated to have been 628-1078 species,[176] as well as many groups
of birds did suddenly become extinct approximately 66 million years ago. It has
been suggested that because small mammals, squamata and birds occupied the
ecological niches suited for small body size, non-avian dinosaurs never evolved
a diverse fauna of small-bodied species, which led to their downfall when large-
bodied terrestrial tetrapods were hit by the mass extinction event.[177] Many other
groups of animals also became extinct at this time,
including ammonites (nautilus-
like mollusks), mosasaurs, plesiosaurs, pterosaurs, and many groups
of mammals.[12] Significantly, the insects suffered no discernible population loss,
which left them available as food for other survivors. This mass extinction is
known as the Cretaceous–Paleogene extinction event. The nature of the event
that caused this mass extinction has been extensively studied since the 1970s;
at present, several related theories are supported by paleontologists. Though
the consensus is that an impact event was the primary cause of dinosaur
extinction, some scientists cite other possible causes, or support the idea that a
confluence of several factors was responsible for the sudden disappearance of
dinosaurs from the fossil record.[178][179][180]

Impact event
Main article: Chicxulub crater

The Chicxulub Crater at the tip of the Yucatán Peninsula; the impactor that formed
this crater may have caused the dinosaur extinction.

The asteroid collision theory, which was brought to wide attention in 1980
by Walter Alvarez and colleagues, links the extinction event at the end of the
Cretaceous period to a bolide impact approximately 66 million years
ago.[181] Alvarez et al. proposed that a sudden increase in iridium levels, recorded
around the world in the period's rock stratum, was direct evidence of the
impact.[182] The bulk of the evidence now suggests that a bolide 5 to 15
kilometers (3.1 to 9.3 miles) wide hit in the vicinity of the Yucatán Peninsula (in
southeastern Mexico), creating the approximately 180 km (110 mi) Chicxulub
Crater and triggering the mass extinction.[183][184] Scientists are not certain
whether dinosaurs were thriving or declining before the impact event. Some
scientists propose that the meteorite impact caused a long and unnatural drop in
Earth's atmospheric temperature, while others claim that it would have instead
created an unusual heat wave. The consensus among scientists who support
this theory is that the impact caused extinctions both directly (by heat from the
meteorite impact) and also indirectly (via a worldwide cooling brought about
when matter ejected from the impact crater reflected thermal radiation from the
sun). Although the speed of extinction cannot be deduced from the fossil record
alone, various models suggest that the extinction was extremely rapid, being
down to hours rather than years.[185]
Deccan Traps
Main article: Deccan Traps
Before 2000, arguments that the Deccan Traps flood basalts caused
the extinction were usually linked to the view that the extinction was gradual, as
the flood basalt events were thought to have started around 68 million years
ago and lasted for over 2 million years. However, there is evidence that two
thirds of the Deccan Traps were created in only 1 million years about 66 million
years ago, and so these eruptions would have caused a fairly rapid extinction,
possibly over a period of thousands of years, but still longer than would be
expected from a single impact event.[186][187]
The Deccan Traps in India could have caused extinction through several
mechanisms, including the release into the air of dust and sulfuric aerosols,
which might have blocked sunlight and thereby reduced photosynthesis in
plants. In addition, Deccan Trap volcanism might have resulted in carbon
dioxide emissions, which would have increased the greenhouse effect when the
dust and aerosols cleared from the atmosphere.[187] Before the mass extinction of
the dinosaurs, the release of volcanic gases during the formation of the Deccan
Traps "contributed to an apparently massive global warming. Some data point to
an average rise in temperature of 8 °C (14 °F) in the last half million years
before the impact [at Chicxulub]."[186][187]
In the years when the Deccan Traps hypothesis was linked to a slower
extinction, Luis Alvarez (who died in 1988) replied that paleontologists were
being misled by sparse data. While his assertion was not initially well-received,
later intensive field studies of fossil beds lent weight to his claim. Eventually,
most paleontologists began to accept the idea that the mass extinctions at the
end of the Cretaceous were largely or at least partly due to a massive Earth
impact. However, even Walter Alvarez has acknowledged that there were other
major changes on Earth even before the impact, such as a drop in sea level and
massive volcanic eruptions that produced the Indian Deccan Traps, and these
may have contributed to the extinctions.[188]

Possible Paleocene survivors

Main article: Paleocene dinosaurs
Non-avian dinosaur remains are occasionally found above the Cretaceous–
Paleogene boundary. In 2001, paleontologists Zielinski and Budahn reported the
discovery of a single hadrosaurleg-bone fossil in the San Juan Basin, New
Mexico, and described it as evidence of Paleocene dinosaurs. The formation in
which the bone was discovered has been dated to the early Paleocene epoch,
approximately 64.5 million years ago. If the bone was not re-deposited into
that stratum by weathering action, it would provide evidence that some dinosaur
populations may have survived at least a half million years into the Cenozoic
Era.[189] Other evidence includes the finding of dinosaur remains in the Hell Creek
Formation up to 1.3 m (51 in) above the Cretaceous–Paleogene boundary,
representing 40000 years of elapsed time. Similar reports have come from other
parts of the world, including China.[190] Many scientists, however, dismissed the
supposed Paleocene dinosaurs as re-worked, that is, washed out of their
original locations and then re-buried in much later sediments.[191][192] Direct dating
of the bones themselves has supported the later date, with U–Pb dating
methods resulting in a precise age of 64.8 ± 0.9 million years ago. [193] If correct,
the presence of a handful of dinosaurs in the early Paleocene would not change
the underlying facts of the extinction.[191]

History of study
Further information: History of paleontology
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, Hua Yang Guo Zhi, a book written
by Chang Qu during the Western Jin Dynasty (265–316), reported the discovery
of dragon bones at Wucheng in Sichuan Province.[194] Villagers in central China
have long unearthed fossilized "dragon bones" for use in traditional medicines, a
practice that continues today.[195] In Europe, dinosaur fossils were generally
believed to be the remains of giants and other biblical creatures.[196]
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,[197] 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 Natural
History of Oxfordshire in 1677. 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 thigh bone of a
giant human similar to those mentioned in the Bible. In 1699, Edward Lhuyd, a
friend of Sir Isaac Newton, was responsible for the first published scientific
treatment of what would now be recognized as a dinosaur when he described
and named a sauropod tooth, "Rutellum implicatum",[198][199] that had been found
in Caswell, near Witney, Oxfordshire.[200]

William Buckland

Between 1815 and 1824, the Rev William Buckland, a professor of geology at
Oxford, collected more fossilized bones of Megalosaurus and became the first
person to describe a dinosaur in a scientific journal.[197][201] The second 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.[202][203]
The study of these "great fossil lizards" soon became of great interest to
European and American scientists, and in 1842 the English
paleontologist Richard Owencoined the term "dinosaur". He 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.[204]
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 dinosaur mania in the United States.[205]

Edward Drinker Cope

Othniel Charles Marsh

Marsh's 1896 illustration of the bones of Stegosaurus, a dinosaur he described and

named in 1877

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. The feud probably
originated when Marsh publicly pointed out that Cope's reconstruction of
an Elasmosaurus skeleton was flawed: Cope had inadvertently placed
the plesiosaur's head at what should have been the animal's tail end. The 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. Marsh 'won' the
contest primarily because he was better funded through a relationship with
the US Geological Survey. 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 (a method modern
paleontologists would find appalling). 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, while Marsh's is on display at the Peabody Museum of Natural
History at Yale University.[206]
After 1897, the search for dinosaur fossils extended to every continent,
including Antarctica. The first Antarctic dinosaur to be discovered,
the ankylosaurid Antarctopelta oliveroi, was found on James Ross Island in
1986,[207] although it was 1994 before an Antarctic species, the
theropod Cryolophosaurus ellioti, was formally named and described in a
scientific journal.[208]
Current dinosaur "hot spots" include southern South America
(especially Argentina) and China. China in particular has produced many
exceptional feathered dinosaur specimens due to the unique geology of its
dinosaur beds, as well as an ancient arid climate particularly conducive to
fossilization.[160]

"Dinosaur renaissance"
Main article: Dinosaur renaissance

Paleontologist Robert T. Bakkerwith mounted skeleton of a tyrannosaurid

(Gorgosaurus libratus)

The field of dinosaur research has enjoyed a surge in activity that began in the
1970s and is ongoing. This was triggered, in part, by John Ostrom's discovery
of Deinonychus, an active predator that may have been warm-blooded, in
marked contrast to the then-prevailing image of dinosaurs as sluggish and cold-
blooded. Vertebrate paleontology has become a global science. Major new
dinosaur discoveries have been made by paleontologists working in previously
unexploited regions, including India, South America, Madagascar, Antarctica,
and most significantly China (the amazingly well-preserved feathered
dinosaurs[5] in China have further consolidated the link between dinosaurs and
their living descendants, modern birds). The widespread application of cladistics,
which rigorously analyzes the relationships between biological organisms, has
also proved tremendously useful in classifying dinosaurs. Cladistic analysis,
among other modern techniques, helps to compensate for an often incomplete
and fragmentary fossil record.[209]

showTimeline of notable dinosaur taxonomic descriptions

Soft tissue and DNA

 Scipionyx from the Natural History Museum of Milan, Italy

One of the best examples of soft-tissue impressions in a fossil dinosaur was

discovered in Pietraroia, Italy. The discovery was reported in 1998, and
described the specimen of a small, very young coelurosaur, Scipionyx
samniticus. The fossil includes portions of the intestines, colon, liver, muscles,
and windpipe of this immature dinosaur.[68]
In the March 2005 issue of Science, the paleontologist Mary Higby
Schweitzer and her team announced the discovery of flexible material
resembling actual soft tissue inside a 68-million-year-old Tyrannosaurus rex leg
bone from the Hell Creek Formation in Montana. After recovery, the tissue was
rehydrated by the science team.[69] When the fossilized bone was treated over
several weeks to remove mineral content from the fossilized bone-marrow cavity
(a process called demineralization), Schweitzer found evidence of intact
structures such as blood vessels, bone matrix, and connective tissue (bone
fibers). Scrutiny under the microscope further revealed that the putative dinosaur
soft tissue had retained fine structures (microstructures) even at the cellular
level. The exact nature and composition of this material, and the implications of
Schweitzer's discovery, are not yet clear.[69]
In 2009, a team including Schweitzer announced that, using even more careful
methodology, they had duplicated their results by finding similar soft tissue in
a duck-billed dinosaur, Brachylophosaurus canadensis, found in the Judith River
Formation of Montana. This included even more detailed tissue, down to
preserved bone cells that seem even to have visible remnants of nuclei and
what seem to be red blood cells. Among other materials found in the bone
was collagen, as in the Tyrannosaurus bone. The type of collagen an animal has
in its bones varies according to its DNA and, in both cases, this collagen was of
the same type found in modern chickens and ostriches.[210]
The extraction of ancient DNA from dinosaur fossils has been reported on two
separate occasions;[211] upon further inspection and peer review, however,
neither of these reports could be confirmed.[212] However, a
functional peptide involved in the vision of a theoretical dinosaur has been
inferred using analytical phylogenetic reconstruction methods on gene
sequences of related modern species such as reptiles and birds.[213] In addition,
several proteins, including hemoglobin,[214] have putatively been detected in
dinosaur fossils.[215][216]
In 2015, researchers reported finding structures similar to blood cells and
collagen fibers, preserved in the bone fossils of six Cretaceous dinosaur
specimens, which are approximately 75 million years old.[217][218]
 Outdated Iguanodon statues created by Benjamin Waterhouse Hawkins for the
Crystal Palace Park in 1853

The battles that may have occurred between Tyrannosaurus rex and Triceratops are
a recurring theme in popular science and dinosaurs' depiction in culture.

Cultural depictions
Main article: Cultural depictions of dinosaurs
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. 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.[219]
Public enthusiasm for dinosaurs first developed in Victorian England, where in
1854, three decades after the first scientific descriptions of dinosaur remains, a
menagerie of lifelike dinosaur sculptures were unveiled in London's Crystal
Palace Park. The Crystal Palace dinosaurs proved so popular that a strong
market in 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.[220] Dinosaurs' enduring popularity, 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.[221]
The popular preoccupation with dinosaurs has ensured their appearance
in literature, film, and other media. Beginning in 1852 with a passing mention
in Charles Dickens' Bleak House,[222] dinosaurs have been featured in large
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 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.[223]

See also
 Animal track
 Dinosaur diet and feeding
 Evolutionary history of life
 Lists of dinosaur-bearing stratigraphic units
 List of dinosaur genera
 List of unavailable dinosaur genera

Notes
1. ^ Although dinosaurs are reptiles, views on them as being cold-
blooded animals have changed in recent years; see physiology for more details.

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Further reading
Library resources about
Dinosaurs

Online books
Resources in your library
Resources in other libraries

 Bakker, Robert T. (1986). The Dinosaur Heresies: New Theories Unlocking

the Mystery of the Dinosaurs and Their Extinction. New York:
Morrow. ISBN 0-688-04287-2.
 Holtz, Thomas R. Jr. (2007). Dinosaurs: The Most Complete, Up-to-Date
Encyclopedia for Dinosaur Lovers of All Ages. New York: Random
House. ISBN 978-0-375-82419-7.
 Paul, Gregory S. (2000). The Scientific American Book of Dinosaurs. New
York: St. Martin's Press. ISBN 0-312-26226-4.
 Paul, Gregory S. (2002). Dinosaurs of the Air: The Evolution and Loss of
Flight in Dinosaurs and Birds. Baltimore: The Johns Hopkins University
Press. ISBN 0-8018-6763-0.
 Randall, Lisa (2015), Dark matter and the dinosaurs: The astounding
interconnectedness of the universe, New York: Harper Collins
Publishers, ISBN 978-0-06-232847-2
 Sternberg, C.M. (1966). Canadian Dinosaurs, in Geological Series, no. 54.
Second ed. [Ottawa]: National Museum of Canada. 28 p., amply ill.
 Stewart, Tabori & Chang (1997). The Humongous Book of Dinosaurs. New
York: Stewart, Tabori & Chang. ISBN 978-1-55670-596-0. Retrieved May
17, 2012. ISBN 1-55670-596-4(Article: The Humongous Book of Dinosaurs)
 Zhou, Z. (2004). "The origin and early evolution of birds: discoveries,
disputes, and perspectives from fossil
evidence" (PDF). Naturwissenschaften. 91 (10): 455–
471. Bibcode:2004NW.....91..455Z. doi:10.1007/s00114-004-0570-
4. PMID 15365634. Archived from the original (PDF) on 2013-05-29.