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Dinosaur
From Wikipedia, the free encyclopedia

Dinosaurs are a diverse group of reptiles[note 1] of the clade Dinosauria that first appeared during the Triassic
Flowers
Mammals Dinosaurs
period. Although
Dinosaursthe exact origin and timing of the evolution of dinosaurs is the subject of active research,[1] the
Land life
current scientific consensus places their origin between 231 and 243 million years ago.[2] They became the Temporal range:
dominant terrestrial vertebrates after the TriassicJurassic extinction event 201 million years ago. Their dominance Late TriassicPresent, 231.4 0 Mya
continued through the Jurassic and Cretaceous periods and ended when the CretaceousPaleogene extinction event
Pre OS D C P T J K Pg N
Multicellular
led to the extinction of most dinosaur groups 66 million years ago.
life
The record indicates that birds are modern feathered dinosaurs,[3] having evolved from theropod ancestors
during the Jurassic Period.[4] As such, birds were the only dinosaur lineage to survive the mass extinction event.[5]
Throughout the remainder of this article, the term "dinosaur" is sometimes used generically to refer to the
combined group of avian dinosaurs (birds) and non-avian dinosaurs (all other dinosaurs); at other times it is used to
refer to the non-avian dinosaurs specifically, while the avian dinosaurs are sometimes simply referred to as "birds".
Eukaryotes
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, are the most diverse group of vertebrates besides perciform fish.[7] Using fossil evidence,
paleontologists have identified over 500 distinct genera[8] 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.[10]
Through the first half of the 20th century, before birds were recognized to be dinosaurs, most of the scientific
photosynthesis
community believed dinosaurs to have been sluggish and cold-blooded. Most research conducted since the 1970s,
A collection of fossil dinosaur skeletons.
however, has indicated that all dinosaurs were active animals with elevated metabolisms and numerous adaptations
for social interaction. Some are herbivorous, others carnivorous. Evidence suggests that egg laying and nest Clockwise from top left: Microraptor gui (a
building are additional traits shared by all dinosaurs. winged theropod), Apatosaurus louisae (a giant
sauropod), Edmontosaurus regalis (a duck-billed
While dinosaurs were ancestrally bipedal, many extinct groups included quadrupedal species, and some were able
Single-celled ornithopod), Triceratops horridus (a horned
to shift between these stances. Elaborate display structures such as horns or crests are common to all dinosaur
groups, and life
some extinct groups developed skeletal modifications such as bony armor and spines. While the ceratopsian), Stegosaurus stenops (a plated
dinosaurs' modern-day surviving avian lineage (birds) are generally small due to the constraints of flight, many stegosaur), Pinacosaurus grangeri (an armored
prehistoric dinosaurs
water (non-avian and avian) were large-bodiedthe largest sauropod dinosaurs are estimated to ankylosaur)
have reached lengths of 39.7 meters (130 feet)[11] and heights of 18 meters (59 feet)[12] 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 Scientific classification
on preservation bias, as large, sturdy bones are more likely to last until they are fossilized. Many dinosaurs were Kingdom: Animalia
quite small: Xixianykus, for example, was only about 50 cm (20 in) long.
Phylum: Chordata
Since the first dinosaur fossils were recognized in the early 19th century, mounted fossil dinosaur skeletons have
Clade: Dinosauriformes
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 Clade: Dinosauria
dinosaurs' regular appearance in best-selling books and films, such as Jurassic Park. Persistent public enthusiasm Owen, 1842
for the animals has resulted in significant funding for dinosaur science, and new discoveries are regularly covered
by the media. Major groups

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

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12 References
13 Further reading
14 External links

Etymology
The taxon Dinosauria was formally named in 1842 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.[13] The term is derived from the Greek words (deinos, meaning "terrible", "potent", or
"fearfully great") and (sauros, meaning "lizard" or "reptile").[13][14] 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.[15]

Other prehistoric animals, including mosasaurs, ichthyosaurs, pterosaurs, plesiosaurs, and Dimetrodon, while often popularly conceived of as dinosaurs, are not
taxonomically classified as dinosaurs.

Definition
Under phylogenetic nomenclature, dinosaurs are usually defined as the group consisting of Triceratops, Neornithes, their
most recent common ancestor (MRCA), and all descendants.[16] 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.[17] 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).[18]

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
Triceratops skeleton, Natural History
paleontologists concerned with dinosaurs reject the traditional style of classification in favor of phylogenetic taxonomy;
Museum of Los Angeles County
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. Birds are classified as
belonging to the subgroup Maniraptora, which are coelurosaurs, which are theropods, which are saurischians, which are
dinosaurs.[19]

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 Ornithoscelida to refer to the group containing Ornithischia and Theropoda. Dinosauria itself was re-defined to 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.[20][21]

General description

Using one of the above definitions, dinosaurs can be generally described as archosaurs with hind limbs held erect
beneath the body.[22] 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.[23] 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.[24]

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.[8] An earlier study
predicted that about 3400 dinosaur genera existed, including many that would not have been preserved in the fossil In phylogenetic taxonomy, birds are included in the
record.[25] By September 17, 2008, 1047 different species of dinosaurs had been named.[9] group Dinosauria.

In 2016, the estimated number of dinosaur species that existed in the Mesozoic era was estimated to be
1,5432,468.[26][27] Some are herbivorous, others carnivorous, including seed-eaters, fish-eaters, insectivores, and omnivores. While dinosaurs were ancestrally
bipedal (as are all modern birds), some prehistoric species were quadrupeds, and others, such as Ammosaurus 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.[10] 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 semi-aquatic habits.[28]

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.[29]

A detailed assessment of archosaur interrelations by Sterling Nesbitt[30] 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

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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.[16]

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 snout and lower jaw.[31] 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);[16] and a perforate acetabulum, or hip socket, with a hole at the center of its inside surface (closed in
Saturnalia, for example).[32][33] 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.[34]

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.[35] 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.[36] Their erect posture
enabled early dinosaurs to breathe easily while moving, which likely permitted stamina and activity levels that surpassed
those of "sprawling" reptiles.[37] Erect limbs probably also helped support the evolution of large size by reducing bending
stresses on limbs.[38] Some non-dinosaurian archosaurs, including rauisuchians, also had erect limbs but achieved this by a
Hip joints and hindlimb postures of:
"pillar erect" configuration of the hip joint, where instead of having a projection from the femur insert on a socket on the hip,
(left to right) typical reptiles
the upper pelvic bone was rotated to form an overhanging shelf.[38]
(sprawling), dinosaurs and mammals
(erect), and rauisuchians (erect)
Evolutionary history
Origins and early evolution
Life timeline
Dinosaurs diverged from their archosaur ancestors during the middle to late Triassic period, roughly view discuss
20 million years after the PermianTriassic extinction event wiped out an estimated 95% of all life on
0 Earliest humans
Earth.[39][40] Radiometric dating of the rock formation that contained fossils from the early dinosaur Quaternary P
h
genus Eoraptor at 231.4 million years old establishes its presence in the fossil record at this time.[41] Karoo
a
n
Paleontologists think that Eoraptor resembles the common ancestor of all dinosaurs;[42] if this is true, its Andean e
-500 r Cambrian explosion
traits suggest that the first dinosaurs were small, bipedal predators.[43] The discovery of primitive, o Ediacara biota
Cryogenian
dinosaur-like ornithodirans such as Marasuchus and Lagerpeton in Argentinian Middle Triassic strata z
o
supports this view; analysis of recovered fossils suggests that these animals were indeed small, bipedal i
c
predators. Dinosaurs may have appeared as early as 243 million years ago, as evidenced by remains of -1000
the genus Nyasasaurus from that period, though known fossils of these animals are too fragmentary to Earliest sexual

tell if they are dinosaurs or very close dinosaurian relatives.[44] P
reproduction
r
-1500 o
When dinosaurs appeared, they were not the dominant terrestrial animals. The terrestrial habitats were t
e
occupied by various types of archosauromorphs and therapsids, like cynodonts and rhynchosaurs. Their r
main competitors were the pseudosuchia, such as aetosaurs, ornithosuchids and rauisuchians, which o
-2000 z
were more successful than the dinosaurs.[45] Most of these other animals became extinct in the Triassic, o
i
in one of two events. First, at about 215 million years ago, a variety of basal archosauromorphs, Huronian c
Oxygen crisis
including the protorosaurs, became extinct. This was followed by the TriassicJurassic extinction event
(about 200 million years ago), that saw the end of most of the other groups of early archosaurs, like -2500 Atmospheric oxygen
aetosaurs, ornithosuchids, phytosaurs, and rauisuchians. Rhynchosaurs and dicynodonts survived (at
least in some areas) at least as late as early-mid Norian and early Rhaetian, respectively,[46][47] and the Pongola

exact date of their extinction is uncertain. These losses left behind a land fauna of crocodylomorphs, -3000 A
r
dinosaurs, mammals, pterosaurians, and turtles.[16] The first few lines of early dinosaurs diversified c
h
through the Carnian and Norian stages of the Triassic, possibly by occupying the niches of the groups e
that became extinct.[18] -3500 na Earliest oxygen

Evolution and paleobiogeography

LHB meteorites
-4000
H
Dinosaur evolution after the Triassic follows changes in vegetation and the location of continents. In the a
late Triassic and early Jurassic, the continents were connected as the single landmass Pangaea, and there d
Earliest life
e
was a worldwide dinosaur fauna mostly composed of coelophysoid carnivores and early a Earliest water
-4500 n Earliest Earth
sauropodomorph herbivores.[48] Gymnosperm plants (particularly conifers), a potential food source,
Axis scale: millions of (4540)
years.
radiated in the late Triassic. Early sauropodomorphs did not have sophisticated mechanisms for
Orange labels: known ice ages.
processing food in the mouth, and so must have employed other means of breaking down food farther
Also see: Human timeline and Nature timeline
along the digestive tract.[49] The general homogeneity of dinosaurian faunas continued into the middle
and late Jurassic, where most localities had predators consisting of ceratosaurians, spinosauroids, and

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carnosaurians, 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 sinraptorid theropods and unusual, long-necked sauropods like
Mamenchisaurus.[48] 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.[49] Another notable evolutionary event of the Jurassic
was the appearance of true birds, descended from maniraptoran coelurosaurians.[50]

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.[48] The stegosaurians appear to have gone extinct at
Skeleton of Marasuchus lilloensis, a
some point in the late early Cretaceous or early late Cretaceous. A major change in the early Cretaceous, which would be
dinosaur-like ornithodiran
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.[49] Some sauropods
also evolved tooth batteries, best exemplified by the rebbachisaurid Nigersaurus.[51]

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.[48] Flowering plants were greatly radiating,[49] with the first
grasses appearing by the end of the Cretaceous.[52] 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.[49] The early forms Herrerasaurus (large),
Eoraptor (small) and a Plateosaurus
The CretaceousPaleogene extinction event, which occurred approximately 66 million years ago at the end of the Cretaceous skull
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.[53]

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 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.[54] Terror birds and bathornithids occupied carnivorous guilds alongside predatory mammals,[55][56] and ratites are still being 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.[57]

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.[29]

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 ()
meaning "lizard" and ischion () meaning "hip joint"retained the hip structure of their ancestors, with a pubis bone directed cranially, or forward.[36] This
basic form was modified by rotating the pubis backward to varying degrees in several groups (Herrerasaurus,[58] therizinosauroids,[59] dromaeosaurids,[60] and
birds[50]). Saurischia includes the theropods (exclusively bipedal and with a wide variety of diets) and sauropodomorphs (long-necked herbivores which include
advanced, quadrupedal groups).[61][62]

By contrast, ornithischians"bird-hipped", from the Greek ornitheios () meaning "of a bird" and ischion () 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".)[29]

Saurischian pelvis structure (left side) Tyrannosaurus pelvis (showing Ornithischian pelvis structure (left side)
saurischian structure left side)

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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).[5] 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)

Theropoda (all bipedal; most were carnivorous)

Herrerasauria (early bipedal carnivores)

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 semi-aquatic spinosaurids) Artist's impression of six dromaeosaurid
Carnosauria (Allosaurus and close relatives, like Carcharodontosaurus) theropods: from left to right
Coelurosauria (feathered theropods, with a range of body sizes and niches)[3] Microraptor, Dromaeosaurus,
Compsognathidae (common early coelurosaurs with reduced forelimbs) Austroraptor, Velociraptor, Utahraptor,
Tyrannosauridae (Tyrannosaurus and close relatives; had reduced forelimbs) and Deinonychus
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)

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") Artist's impression of four macronarian
Cetiosauridae ("whale reptiles") sauropods: from left to right
Turiasauria (European group of Jurassic and Cretaceous sauropods) Camarasaurus, Brachiosaurus,
Neosauropoda ("new sauropods") Giraffatitan, and Euhelopus
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)

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)

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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)
Restoration of six ornithopods; far left:
Biology Camptosaurus, left: Iguanodon, center
background: Shantungosaurus, center
foreground: Dryosaurus, right:
Knowledge about dinosaurs is derived from a variety of fossil and non-fossil records, including fossilized bones, feces,
Corythosaurus, far right (large)
trackways, gastroliths, feathers, impressions of skin, internal organs and soft tissues.[63][64] Many fields of study contribute to
Tenontosaurus.
our understanding of dinosaurs, including physics (especially biomechanics), chemistry, biology, and the earth sciences (of
which paleontology is a sub-discipline).[65][66] Two topics of particular interest and study have been dinosaur size and
behavior.[67]

Size

Current evidence suggests that dinosaur average size varied through the Triassic, early Jurassic,
late Jurassic and Cretaceous periods.[42] Predatory theropod dinosaurs, which occupied most
terrestrial carnivore niches during the Mesozoic, most often fall into the 100 to 1 000 kg (220 to
2 200 lb) category when sorted by estimated weight into categories based on order of magnitude,
whereas recent predatory carnivoran mammals peak in the 10 to 100 kg (22 to 220 lb)
category.[68] The mode of Mesozoic dinosaur body masses is between one and ten metric
Scale diagram comparing the average human to the largest known
tonnes.[69] This contrasts sharply with the size of Cenozoic mammals, estimated by the National dinosaurs in five major clades: Sauropoda (Argentinosaurus
Museum of Natural History as about 2 to 5 kg (4.4 to 11.0 lb).[70] huinculensis), Ornithopoda (Shantungosaurus giganteus), Theropoda
(Spinosaurus aegyptiacus), Thyreophora (Stegosaurus armatus) and
The sauropods were the largest and heaviest dinosaurs. For much of the dinosaur era, the smallest
Marginocephalia (Triceratops prorsus)
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.[71] 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.[12]

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.[72]

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 fr Naturkunde Berlin;[73] this
mount is 12 meters (39 ft) tall and 21.822.5 meters (7274 ft) long,[74][75] and would have belonged to an animal that
weighed between 30 000 and 60 000 kilograms (70 000 and 130 000 lb). The longest complete dinosaur is the 27 meters (89
feet) long Diplodocus, which was discovered in Wyoming in the United States and displayed in Pittsburgh's Carnegie Natural
History Museum in 1907.[76]
Comparative size of Giraffatitan to the
There were larger dinosaurs, but knowledge of them is based entirely on a small average human
number of fragmentary fossils. Most of the largest herbivorous specimens on record
were discovered in the 1970s or later, and include the massive Argentinosaurus,
which may have weighed 80 000 to 100 000 kilograms (90 to 110 short tons); some of the longest were the 33.5 meters
(110 ft) long Diplodocus hallorum[12] (formerly Seismosaurus) and the 3334 meters (108112 ft) long Supersaurus;[77] 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
122 400 kg (270 000 lb).[12] 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.[78] The largest known
carnivorous dinosaur was Spinosaurus, reaching a length of 12.6 to 18 meters (41 to 59 ft), and weighing 720.9 tonnes
Comparative size of Eoraptor to the
(7.723 short tons).[79][80] Other large carnivorous theropods included Giganotosaurus, Carcharodontosaurus and
average human
Tyrannosaurus.[80] Therizinosaurus and Deinocheirus were among the tallest of the theropods.

The smallest dinosaur known is the bee hummingbird,[81] with a length of only 5 cm (2.0 in) and mass of around 1.8 g
(0.063 oz).[82]The smallest known non-avialan dinosaurs were about the size of pigeons and were those theropods most closely related to birds.[83] For example,
Anchiornis huxleyi is currently the smallest non-avialan dinosaur described from an adult specimen, with an estimated weight of 110 grams[84] and a total skeletal
length of 34 cm (1.12 ft).[83][84] The smallest herbivorous non-avialan dinosaurs included Microceratus and Wannanosaurus, at about 60 cm (2.0 ft) long each.[5][85]

Behavior

Many modern birds are highly social, often found living in flocks. There is general agreement that some behaviors that are common in birds, as well as in crocodiles
(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.[65]

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.[86] 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

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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,[87] although there is no evidence for specific herd structures.[88]
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.[89] The interpretation of dinosaurs as gregarious has also extended to depicting carnivorous theropods as
pack hunters working together to bring down large prey.[90][91] 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 A nesting ground of hadrosaur
many modern diapsid predators.[92] Maiasaura peeblesorum was discovered
in 1978.
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.[93]

Artist's rendering of two Centrosaurus
apertus engaged in intra-specific
combat
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,[94] providing evidence that dinosaurs did indeed attack each other.[95]
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.[95] Cannibalism amongst some
species of dinosaurs was confirmed by tooth marks found in Madagascar in 2003, involving the theropod Majungasaurus.[96]

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.[97]

Based on current fossil evidence from dinosaurs such as Oryctodromeus, some ornithischian species seem to have led a partially fossorial (burrowing) lifestyle.[98]
Many modern birds are arboreal (tree climbing), and this was also true of many Mesozoic birds, especially the enantiornithines.[99] While some early bird-like
species may have already been arboreal as well (including dromaeosaurids such as Microraptor[100]) 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,[101] whether diplodocids could create sonic booms via whip-like tail snapping,[102] and whether sauropods could
float.[103]

Communication

Modern birds are known to communicate using visual and auditory signals, and the wide diversity of visual display structures
among fossil dinosaur groups, such as horns, frills, crests, sails and feathers, suggests that visual communication has always
been important in dinosaur biology.[104] Reconstruction of the plumage color of Anchiornis huxleyi, suggest the importance
of color in visual communication in non-avian dinosaurs.[105] 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.[104]

The earliest remains of a syrinx, which has enough mineral content for fossilization, was found in a specimen of the duck-
Artist's impression of a striking and
like Vegavis iaai dated 69-66 million year ago, and this organ is unlikely to have existed in non-avian dinosaurs. However, in unusual visual display in a
contrast to Senter, the researchers have suggested that dinosaurs could vocalize and that the syrinx-based vocal system of Lambeosaurus magnicristatus
birds evolved from a larynx-based one, rather than the two systems evolving independently. [106] 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.[107] The crests of the Lambeosaurini and
nasal chambers of ankylosaurids have been suggested to function in vocal resonance,[108][109] 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.[104]

Reproductive biology

All dinosaurs lay amniotic eggs with hard shells made mostly of calcium carbonate.[110] Eggs are usually laid in a nest. Most
species create somewhat elaborate nests, which can be cups, domes, plates, beds scrapes, mounds, or burrows.[111] 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.[112]

Nest of a plover (Charadrius) 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.[113]

Another widespread trait among modern birds 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, suggesting this behavior might also have been common to all
dinosaurs.[114] There is evidence that other non-theropod dinosaurs, like Patagonian titanosaurian sauropods, also nested in large groups.[115] A specimen of the
Mongolian oviraptorid Citipati osmolskae was discovered in a chicken-like brooding position in 1993,[116] which may indicate that they had begun using an

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insulating layer of feathers to keep the eggs warm.[117] Parental care being a trait common to all dinosaurs is supported by
other finds. For example, a dinosaur embryo (pertaining to the prosauropod Massospondylus) was found without teeth,
indicating that some parental care was required to feed the young dinosaurs.[118] Trackways have also confirmed parental
behavior among ornithopods from the Isle of Skye in northwestern Scotland.[119] Nests and eggs have been found for most
major groups of dinosaurs, and it appears likely that all dinosaurs cared for their young to some extent either before or shortly
after hatching.[120]

Physiology
Fossil interpreted as a nesting
Because both modern crocodilians and birds have four-chambered hearts (albeit modified in crocodilians), it is likely that this oviraptorid Citipati at the American
is a trait shared by all archosaurs, including all dinosaurs.[121] While all modern birds have high metabolisms and are "warm Museum of Natural History. Smaller
blooded" (endothermic), a vigorous debate has been ongoing since the 1960s regarding how far back in the dinosaur lineage fossil far right showing inside one of the
this trait extends. Scientists disagree as to whether non-avian dinosaurs were endothermic, ectothermic, or some combination eggs.
of both.[122]

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.[123]

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.[124][125]

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.[126] Early avian-style respiratory systems with air sacs may have been capable of sustaining higher activity levels
than mammals of similar size and build could sustain. 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.[127]

Like other reptiles, dinosaurs are primarily uricotelic, that is, their kidneys extract nitrogenous wastes from their bloodstream
and excrete it as uric acid instead of urea or ammonia via the ureters into the intestine. In most living species, uric acid is Comparison between the air sacs of an
excreted along with feces as a semisolid waste.[128][129][130] However, at least some modern birds (such as hummingbirds) abelisaur and a bird
can be facultatively ammonotelic, excreting most of the nitrogenous wastes as ammonia.[131] They also excrete creatine,
rather than creatinine like mammals.[132] This material, as well as the output of the intestines, emerges from the cloaca.
[133][134] 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.[135]

Origin of birds
The possibility that dinosaurs were the ancestors of birds was first suggested in 1868 by Thomas Henry Huxley.[136] 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.[137] However, as later discoveries showed, clavicles (or a single fused wishbone, which derived from
separate clavicles) were not actually absent;[50] they had been found as early as 1924 in Oviraptor, but misidentified as an interclavicle.[138] In the 1970s, John
Ostrom revived the dinosaurbird theory,[139] which gained momentum in the coming decades with the advent of cladistic analysis,[140] and a great increase in the
discovery of small theropods and early birds.[31] 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.[3][50] Birds share over a hundred distinct anatomical features with theropod dinosaurs, which are now generally
accepted to have been their closest ancient relatives.[141] They are most closely allied with maniraptoran coelurosaurs.[50] 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,[142] or that maniraptoran
theropods are the ancestors of birds but themselves are not dinosaurs, only convergent with dinosaurs.[143]

Feathers

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.[144] Direct fossil evidence of feathers or feather-like
structures has been discovered in a diverse array of species in many non-avian dinosaur groups,[3] both among saurischians
and ornithischians. Simple, branched, feather-like structures are known from heterodontosaurids, primitive
neornithischians[145] and theropods,[146] 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.[50][147] Feather-like structures known as pycnofibres have also been found in
pterosaurs,[148] suggesting the possibility that feather-like filaments may have been common in the bird lineage and evolved
before the appearance of dinosaurs themselves.[144] 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.[149]
Various feathered non-avian dinosaurs,
Archaeopteryx was the first fossil found that revealed a potential connection between dinosaurs and birds. It is considered a including Archaeopteryx, Anchiornis,
transitional fossil, in that it displays features of both groups. Brought to light just two years after Darwin's seminal The Microraptor and Zhenyuanlong
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.[150] 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 lagersttte 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.[151]

The description of feathered dinosaurs has not been without controversy; perhaps the most vocal critics have been Alan Feduccia and Theagarten Lingham-Soliar,

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who have proposed that some purported feather-like fossils are the result of the decomposition of collagenous fiber that underlaid the dinosaurs' skin,[152][153][154]
and that maniraptoran dinosaurs with vaned feathers were not actually dinosaurs, but convergent with dinosaurs.[143][153] 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.[155]

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.[3][156][157]

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.[158]

Soft anatomy

Large meat-eating dinosaurs had a complex system of air sacs similar to those found in modern birds, according to a 2005
investigation led by Patrick M. O'Connor. The lungs of theropod dinosaurs (carnivores that walked on two legs and had bird-
like feet) likely pumped air into hollow sacs in their skeletons, as is the case in birds. "What was once formally considered
unique to birds was present in some form in the ancestors of birds", O'Connor said.[159] 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.[160][161]

Behavioral evidence

Fossils of the troodonts Mei and Sinornithoides demonstrate that some dinosaurs slept with their heads tucked under their
arms.[162] 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.[163] 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.[164]
Pneumatopores on the left ilium of
Some dinosaurs are known to have used gizzard stones like modern birds. These stones are swallowed by animals to aid Aerosteon riocoloradensis
digestion and break down food and hard fibers once they enter the stomach. When found in association with fossils, gizzard
stones are called gastroliths.[165]

Extinction of major groups

The discovery that birds are a type of dinosaur showed that dinosaurs in general are not, in fact, extinct as is commonly stated.[166] However, all non-avian dinosaurs
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.[167] Many other groups of animals also became extinct at this time, including
ammonites (nautilus-like mollusks), mosasaurs, plesiosaurs, pterosaurs, and many groups of mammals.[10] Significantly, the insects suffered no discernible
population loss, which left them available as food for other survivors. This mass extinction is known as the CretaceousPaleogene 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.[168][169][170]

Impact event

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.[171] 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.[172] 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 Yucatn Peninsula (in southeastern Mexico), creating the approximately 180 km (110 mi) Chicxulub Crater
and triggering the mass extinction.[173][174] 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.[175]

Deccan Traps The Chicxulub Crater at the tip of the

Yucatn Peninsula; the impactor that
Before 2000, arguments that the Deccan Traps flood basalts caused the extinction were usually linked to the view that the formed this crater may have caused the
extinction was gradual, as the flood basalt events were thought to have started around 68 million years ago and lasted for dinosaur extinction.
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.[176][177]

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.[177] 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]."[176][177]

In the years when the Deccan Traps theory 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

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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.[178]

Possible Paleocene survivors

Non-avian dinosaur remains are occasionally found above the CretaceousPaleogene boundary. In 2001, paleontologists Zielinski and Budahn reported the discovery
of a single hadrosaur leg-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.[179] Other evidence includes
the finding of dinosaur remains in the Hell Creek Formation up to 1.3 m (51 in) above the CretaceousPaleogene boundary, representing 40 000 years of elapsed
time. Similar reports have come from other parts of the world, including China.[180] 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.[181][182] Direct dating of the bones themselves has supported the
later date, with UPb dating methods resulting in a precise age of 64.8 0.9 million years ago.[183] If correct, the presence of a handful of dinosaurs in the early
Paleocene would not change the underlying facts of the extinction.[181]

History of study
Dinosaur fossils have been known for millennia, although their true nature was not recognized. The Chinese, whose modern word for dinosaur is knglng (, or
"terrible dragon"), 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 (265316), reported the discovery of dragon bones at Wucheng in Sichuan Province.[184] Villagers in central China have long unearthed
fossilized "dragon bones" for use in traditional medicines, a practice that continues today.[185] In Europe, dinosaur fossils were generally believed to be the remains
of giants and other biblical creatures.[186]

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,[187] 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",[188][189] that had been found in Caswell, near Witney, Oxfordshire.[190]

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.[187][191] 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.[192][193]

The study of these "great fossil lizards" soon became of great interest to European and American scientists, and in 1842 the English
paleontologist Richard Owen coined 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
William Buckland national collection of dinosaur fossils and other biological and geological exhibits.[194]

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.[195]

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 Othniel Charles Marsh
Edward Drinker Cope
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.[196]

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,[197] although it was 1994
before an Antarctic species, the theropod Cryolophosaurus ellioti, was formally named and described in a scientific
journal.[198]

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.[151] Marsh's 1896 illustration of the bones of
Stegosaurus, a dinosaur he described
"Dinosaur renaissance" and named in 1877

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[3] 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.[199]

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Timeline of notable dinosaur taxonomic descriptions

Paleontologist Robert T. Bakker with

mounted skeleton of a tyrannosaurid
(Gorgosaurus libratus)

Soft tissue and DNA

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.[63]

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.[64] 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.[64]

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.[200]

The extraction of ancient DNA from dinosaur fossils has been reported on two separate occasions;[201] upon further inspection and peer review, however, neither of
these reports could be confirmed.[202] 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.[203] In addition, several proteins, including hemoglobin,[204] have
putatively been detected in dinosaur fossils.[205][206]

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.[207][208]

Cultural depictions
By human standards, dinosaurs were creatures of fantastic appearance and often enormous size. As such, they have captured
the popular imagination and become an enduring part of human culture. 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.[209]

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. Outdated Iguanodon statues created by
The Crystal Palace dinosaurs proved so popular that a strong market in smaller replicas soon developed. In subsequent Benjamin Waterhouse Hawkins for the
decades, dinosaur exhibits opened at parks and museums around the world, ensuring that successive generations would be Crystal Palace Park in 1853
introduced to the animals in an immersive and exciting way.[210] 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.[211]

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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,[212] 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.[213]
The battles that may have occurred
See also between Tyrannosaurus rex and
Triceratops are a recurring theme in
popular science and dinosaurs' depiction
Animal track Lists of dinosaur-bearing stratigraphic units
in culture.
Dinosaur diet and feeding List of dinosaur genera
Evolutionary history of life Living dinosaur

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
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" (http://www.worldcat.org/title/humongous-book-of-dinosaurs
/oclc/36301321&referer=brief_results). New York: Stewart, Tabori & Chang. 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" (https://web.archive.org
/web/20130529094636/http://www.cisneros-heredia.org/infotrans/usfq/ornitofauna/pdfs/zhou2004.pdf) (PDF). Naturwissenschaften. 91 (10): 455471.
Bibcode:2004NW.....91..455Z (http://adsabs.harvard.edu/abs/2004NW.....91..455Z). PMID 15365634 (https://www.ncbi.nlm.nih.gov/pubmed/15365634).
doi:10.1007/s00114-004-0570-4 (https://doi.org/10.1007%2Fs00114-004-0570-4). Archived from the original (http://www.cisneros-heredia.org/infotrans
/usfq/ornitofauna/pdfs/zhou2004.pdf) (PDF) on 2013-05-29.

External links
General

DinoDatabase.com | Hundreds of dinosaurs and dinosaur related topics (http://www.dinodatabase.com/)

Images

The Science and Art of Gregory S. Paul (http://www.gspauldino.com/) Influential paleontologist's anatomy art and paintings
Skeletal Drawing (http://skeletaldrawing.com/) Professional restorations of numerous dinosaurs, and discussions of dinosaur anatomy.
"Dinosaur Discovery" (http://lhldigital.lindahall.org/cdm/landingpage/collection/dino), a collection of images from early works on dinosaurs at the Linda Hall
Library, in support of the exhibition, Paper Dinosaurs, 18241969 (http://dino.lindahall.org/).

Video

BBC Nature: Dinosaur reconstructions and expert interpretations, including Walking with Dinosaurs (http://www.bbc.co.uk/nature/life/dinosaur)
BBC Explainer Dinosaurs a complete history in 4 minutes, animation (http://vimeo.com/74636204)
Origin, evolution and extinction of the dinosaurs (http://www.scientificamerican.com/article/how-dinosaurs-got-their-start-and-met-their-end-video/) Stephen
Brusatte video lecture April 15, 2014

Popular

Dinosaurs & other extinct creatures (http://www.nhm.ac.uk/nature-online/life/dinosaurs-other-extinct-creatures/index.html): From the Natural History
Museum, a well illustrated dinosaur directory.
Dinosaurnews (http://www.dinosaurnews.org/) Dinosaur-related headlines from around the world, including finds and discoveries, and many links.
Dinosauria (http://www.ucmp.berkeley.edu/diapsids/dinosaur.html) From UC Berkeley Museum of Paleontology.
LiveScience.com (http://www.livescience.com/dinosaurs/) Dinosaur pages
Zoom Dinosaurs (http://www.enchantedlearning.com/subjects/dinosaurs/) From Enchanted Learning. Kids' site, info pages and stats, theories, history.
Dinosaur genus list (http://www.geol.umd.edu/~tholtz/dinoappendix/HoltzappendixWinter2010.pdf) contains data tables on nearly every published Mesozoic
dinosaur genus as of January 2011.

Technical

Palaeontologia Electronica (http://palaeo-electronica.org/) From Coquina Press. Online technical journal.

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