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Dinosaur
From Wikipedia, the free encyclopedia
For other uses, see Dinosaur (disambiguation).
Dinosaurs
Temporal range: Late TriassicPresent, 231.4 0 Mya
Pre?
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O
S
D
C
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T
J
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Pg
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Dinosauria montage 2.jpg
A collection of fossil dinosaur skeletons. Clockwise from top left: Microraptor gui
(a winged theropod), Apatosaurus louisae (a giant sauropod), Edmontosaurus regalis
(a duck-billed ornithopod), Triceratops horridus (a horned ceratopsian),
Stegosaurus stenops (a plated stegosaur), Pinacosaurus grangeri (an armored
ankylosaur)
Scientific classification e
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauriformes
Clade: Dinosauria
Owen, 1842
Major groups

Ornithischia
Heterodontosauridae
Genasauria
Neornithischia
Thyreophora
Saurischia
Sauropodomorpha
Theropoda

Dinosaurs are a diverse group of reptiles of the clade Dinosauria that first
appeared during the Triassic period. Although the exact origin and timing of the
evolution of dinosaurs is the subject of active research,[1] the current scientific
consensus places their origin between 231 and 243 million years ago.[2] They became
the dominant terrestrial vertebrates after the TriassicJurassic extinction event
201 million years ago. Their dominance continued through the Jurassic and
Cretaceous periods and ended when the CretaceousPaleogene extinction event led to
the extinction of most dinosaur groups 66 million years ago.

The fossil 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; 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". 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,[6] 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.[9] 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 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 are herbivorous, others carnivorous. Evidence suggests that egg
laying and nest building are additional traits shared by all dinosaurs.

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-bodiedthe largest sauropod dinosaurs are estimated to 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 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

1 Etymology
2 Definition
2.1 General description
2.2 Distinguishing anatomical features
3 Evolutionary history
3.1 Origins and early evolution
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 and references
12 Further reading
13 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 de???? (deinos, meaning "terrible", "potent", or "fearfully
great") and sa???? (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
Triceratops skeleton, Natural History Museum of Los Angeles County

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
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. 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
montage of four birds
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.[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 record.[25] By September 17, 2008,
1047 different species of dinosaurs had been named.[9]

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
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]
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.[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 "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.[38]
Evolutionary history
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Orange labels: known ice ages.
Also see: Human timeline and Nature timeline
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 PermianTriassic extinction
event wiped out an estimated 95% of all life on Earth.[39][40] 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.[41] Paleontologists think that Eoraptor resembles the common ancestor of all
dinosaurs;[42] if this is true, its traits suggest that the first dinosaurs were
small, bipedal predators.[43] 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.[44]

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.[45] 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 TriassicJurassic 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 early Rhaetian, respectively,[46][47] and the exact date of their extinction is
uncertain. These losses left behind a land fauna of crocodylomorphs, dinosaurs,
mammals, pterosaurians, and turtles.[16] 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.[18]
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.[48] 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.[49] 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 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]
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.[48] 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.[49] Some sauropods also evolved tooth
batteries, best exemplified by the rebbachisaurid Nigersaurus.[51]
Full skeleton of an early carnivorous dinosaur, displayed in a glass case in a
museum
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.[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 CretaceousPaleogene 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.[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
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.[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 (sa????)
meaning "lizard" and ischion (?s????) 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 (?????e???)

meaning "of a bird" and ischion (?s????) 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 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).[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)

Artist's impression of six dromaeosaurid theropods: from left to right Microraptor,

Dromaeosaurus, Austroraptor, Velociraptor, Utahraptor, and Deinonychus

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)
Carnosauria (Allosaurus and close relatives, like
Carcharodontosaurus)
Coelurosauria (feathered theropods, with a range of body sizes
and niches)[3]

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)

Artist's impression of six ornithopods and one heterodontosaurid. Far left:

Camptosaurus, left: Iguanodon, center background: Shantungosaurus, center
foreground: Dryosaurus, right: Corythosaurus, far right (small): Heterodontosaurus,
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.[63][64] 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).[65][66] Two topics of particular interest and study have been
dinosaur size and behavior.[67]
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.[42] 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.[68]
The mode of Mesozoic dinosaur body masses is between one and ten metric tonnes.[69]
This contrasts sharply with the size of Cenozoic mammals, estimated by the National
Museum of Natural History as about 2 to 5 kg (4.4 to 11.0 lb).[70]

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.[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]
Comparative size of Giraffatitan 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 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
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.[76]
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); 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 122400 kg (270000 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
(7.723 short tons).[79][80] Other large carnivorous theropods included
Giganotosaurus, Carcharodontosaurus and 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
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.[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 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
many modern diapsid predators.[92]
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.[93]

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

All dinosaurs 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]

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]
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 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
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
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.[121] 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.[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]
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.[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 excreted along with feces as a semisolid waste.[128][129]
[130] However, at least some modern birds (such as hummingbirds) 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
Main article: 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
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.
[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]

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.[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, 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
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.[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]

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.[165]
Extinction of major groups
Main article: CretaceousPaleogene 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.[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
Main article: Chicxulub crater
The Chicxulub Crater at the tip of the Yucatn 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.[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
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.[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
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
Main article: Paleocene dinosaurs

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 40000 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
Further information: History of paleontology

Dinosaur fossils have been known for millennia, although their true nature was not
recognized. The Chinese, whose modern word for dinosaur is konglng (??, 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]
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.[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 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]
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.
[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]
"Dinosaur renaissance"
Main article: Dinosaur renaissance
Paleontologist Robert T. Bakker with 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[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]
[show]Timeline of notable dinosaur taxonomic descriptions
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]
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.[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. 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.[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]

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]
See also

Animal track
Dinosaur diet and feeding
Evolutionary history of life
Lists of dinosaur-bearing stratigraphic units
List of dinosaur genera
Living dinosaur

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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. 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): 455471. Bibcode:2004NW.....91..455Z. PMID 15365634. doi:10.1007/s00114-004-
0570-4. Archived from the original (PDF) on 2013-05-29.

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[hide]

v t e

Basal ornithodirans (Dinosauromorpha and Pterosauromorpha)

Kingdom: Animalia Phylum: Chordata Class: Sauropsida Clade: Archosauria Clade:

Avemetatarsalia
Dinosauromorpha Pterosauromorpha

[hide]
Dinosauromorpha
Lagerpetidae

Dromomeron Lagerpeton Ixalerpeton

Dinosauriformes

Marasuchus Lagosuchus Saltopus Nyasasaurus?

Silesauridae

Lewisuchus Asilisaurus Eucoelophysis Silesaurus Sacisaurus Diodorus

Pseudolagosuchus Lutungutali Ignotosaurus Agnosphitys? Technosaurus?

Dinosauria
(Dinosaurs)

Ornithischia
see below

Saurischia

Basal

Alwalkeria Teyuwasu?

Eusaurischia

Eoraptor Sauropodomorpha
see below Theropoda
see below

Silesaurus opolensis
[show]
Pterosauromorpha
[show]
Ornithischia
[show]
Sauropodomorpha
[show]
Theropoda

Major clades underlined See also categories:

Archosaurs Dinosaurs Pterosaurs Ornithodirans CretaceousPaleogene
extinction event

Taxon identifiers

EoL: 3014672 NCBI: 436486 Fossilworks: 91968

Authority control

LCCN: sh85038094 GND: 4012362-5 BNF: cb124343565 (data) NDL: 00567254

Tyrannoskull.jpgDinosaurs portal Abyssal Brachiopod 00148.jpgPaleontology

portal

Categories:

Carnian first appearancesDinosaursExtant Late Triassic first appearancesFossil

taxa described in 1842Paraphyletic groupsTaxa named by Richard Owen

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