Definition

Under phylogenetic nomenclature, dinosaurs are usually defined as the group consisting of
the most recent common ancestor (MRCA) of Triceratops and modern birds (Neornithes),
and all its descendants.[7] It has also been suggested that Dinosauria be defined with
respect to the MRCA of Megalosaurus and Iguanodon, because these were two of the
three genera cited by Richard Owen when he recognized the Dinosauria.[8] Both definitions
result in the same set of animals being defined as dinosaurs: "Dinosauria
= Ornithischia + Saurischia". This definition includes major groups such
as ankylosaurians (armored herbivorous quadrupeds), stegosaurians (plated herbivorous
quadrupeds), ceratopsians (bipedal or quadrupedal herbivores with neck
frills), pachycephalosaurians (bipedal herbivores with thick skulls), ornithopods (bipedal or
quadrupedal herbivores including "duck-bills"), theropods (mostly bipedal carnivores and
birds), and sauropodomorphs (mostly large herbivorous quadrupeds with long necks and
tails).[9]
Birds are now recognized as being the sole surviving lineage of theropod dinosaurs. In
traditional taxonomy, birds were considered a separate class that had evolved from
dinosaurs, a distinct superorder. However, a majority of contemporary paleontologists
concerned with dinosaurs reject the traditional style of classification in favor
of phylogenetic taxonomy; this approach requires that, for a group to be natural, all
descendants of members of the group must be included in the group as well. Birds are thus
considered to be dinosaurs and dinosaurs are, therefore, not extinct. [10] Birds are classified
as belonging to the subgroup Maniraptora, which are coelurosaurs, which are theropods,
which are saurischians, which are dinosaurs.[11]
Research by Matthew G. Baron, David B. Norman, and Paul M. Barrett in 2017 suggested
a radical revision of dinosaurian systematics. Phylogenetic analysis by Baron et
al. recovered the Ornithischia as being closer to the Theropoda than the
Sauropodomorpha, as opposed to the traditional union of theropods with
sauropodomorphs. They resurrected the clade Ornithoscelida to refer to the group
containing Ornithischia and Theropoda. Dinosauria itself was re-defined as the last
common ancestor of Triceratops horridus, Passer domesticus and Diplodocus carnegii, and
all of its descendants, to ensure that sauropods and kin remain included as dinosaurs. [12][13]

General description

Triceratops skeleton, Natural History Museum of Los Angeles County

Using one of the above definitions, dinosaurs can be generally described

as archosaurs with hind limbs held erect beneath the body.[14] Other prehistoric animals,
including pterosaurs, mosasaurs, ichthyosaurs, plesiosaurs, and Dimetrodon, while often
popularly conceived of as dinosaurs, are not taxonomically classified as
dinosaurs.[15] Pterosaurs are distantly related to dinosaurs, being members of the
clade Ornithodira. The other groups mentioned are, like dinosaurs and pterosaurs,
members of Sauropsida (the reptile and bird clade), except Dimetrodon (which is
a synapsid). None of them had the erect hind limb posture characteristic of true
dinosaurs.[16]
Dinosaurs were the dominant terrestrial vertebrates of the Mesozoic Era, especially the
Jurassic and Cretaceous periods. Other groups of animals were restricted in size and
niches; mammals, for example, rarely exceeded the size of a domestic cat, and were
generally rodent-sized carnivores of small prey.[17] They have always been recognized as
an extremely varied group of animals; over 900 non-avian dinosaur genera have been
identified with certainty as of 2018, and the total number of genera preserved in the fossil
record has been estimated at around 1850, nearly 75% of which remain to be discovered,
and 1124 species by 2016.[18][19][20] A 1995 study predicted that about 3,400 dinosaur genera
ever existed, including many that would not have been preserved in the fossil record. [21]
In 2016, the estimated number of dinosaur species that existed in the Mesozoic was 1,543–
2,468.[22][23] In 2021, the number of modern-day birds (avian dinosaurs) was estimated to be
at 10,806 species.[24] Some are herbivorous, others carnivorous, including seed-eaters, fish-
eaters, insectivores, and omnivores. While dinosaurs were ancestrally bipedal (as are all
modern birds), some prehistoric species were quadrupeds, and others, such
as Anchisaurus and Iguanodon, could walk just as easily on two or four legs. Cranial
modifications like horns and crests are common dinosaurian traits, and some extinct
species had bony armor. Although known for large size, many Mesozoic dinosaurs were
human-sized or smaller, and modern birds are generally small in size. Dinosaurs today
inhabit every continent, and fossils show that they had achieved global distribution by at
least the Early Jurassic epoch.[25] Modern birds inhabit most available habitats, from
terrestrial to marine, and there is evidence that some non-avian dinosaurs (such
as Microraptor) could fly or at least glide, and others, such as spinosaurids,
had semiaquatic habits.[26]

Distinguishing anatomical features

While recent discoveries have made it more difficult to present a universally agreed-upon
list of their distinguishing features, nearly all dinosaurs discovered so far share certain
modifications to the ancestral archosaurian skeleton, or are clearly descendants of older
dinosaurs showing these modifications. Although some later groups of dinosaurs featured
further modified versions of these traits, they are considered typical for Dinosauria; the
earliest dinosaurs had them and passed them on to their descendants. Such modifications,
originating in the most recent common ancestor of a certain taxonomic group, are called
the synapomorphies of such a group.[27]

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

A detailed assessment of archosaur interrelations by Sterling Nesbitt[28] confirmed or found

the following twelve unambiguous synapomorphies, some previously known:

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

fenestra, the main opening in the rear skull roof
 Epipophyses, obliquely backward-pointing processes on the rear top corners of the
anterior (front) neck vertebrae behind the atlas and axis, the first two neck vertebrae
 Apex of a deltopectoral crest (a projection on which the deltopectoral muscles attach)
located at or more than 30% down the length of the humerus (upper arm bone)
 Radius, a lower arm bone, shorter than 80% of humerus length
 Fourth trochanter (projection where the caudofemoralis muscle attaches on the inner
rear shaft) on the femur (thigh bone) is a sharp flange
 Fourth trochanter asymmetrical, with distal, lower, margin forming a steeper angle to
the shaft
 On the astragalus and calcaneum, upper ankle bones, the proximal articular facet, the
top connecting surface, for the fibula occupies less than 30% of the transverse width of
the element
 Exoccipitals (bones at the back of the skull) do not meet along the midline on the floor
of the endocranial cavity, the inner space of the braincase
 In the pelvis, the proximal articular surfaces of the ischium with the ilium and
the pubis are separated by a large concave surface (on the upper side of the ischium a
part of the open hip joint is located between the contacts with the pubic bone and the
ilium)
 Cnemial crest on the tibia (protruding part of the top surface of the shinbone) arcs
anterolaterally (curves to the front and the outer side)
 Distinct proximodistally oriented (vertical) ridge present on the posterior face of the
distal end of the tibia (the rear surface of the lower end of the shinbone)
 Concave articular surface for the fibula of the calcaneum (the top surface of the
calcaneum, where it touches the fibula) has a hollow profile
Nesbitt found a number of further potential synapomorphies and discounted a number of
synapomorphies previously suggested. Some of these are also present in silesaurids,
which Nesbitt recovered as a sister group to Dinosauria, including a large anterior
trochanter, metatarsals II and IV of subequal length, reduced contact between ischium and
pubis, the presence of a cnemial crest on the tibia and of an ascending process on the
astragalus, and many others.[7]

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

A variety of other skeletal features are shared by dinosaurs. However, because they are
either common to other groups of archosaurs or were not present in all early dinosaurs,
these features are not considered to be synapomorphies. For example, as diapsids,
dinosaurs ancestrally had two pairs of Infratemporal fenestrae (openings in the skull behind
the eyes), and as members of the diapsid group Archosauria, had additional openings in
the snout and lower jaw.[29] Additionally, several characteristics once thought to be
synapomorphies are now known to have appeared before dinosaurs, or were absent in the
earliest dinosaurs and independently evolved by different dinosaur groups. These include
an elongated scapula, or shoulder blade; a sacrum composed of three or more fused
vertebrae (three are found in some other archosaurs, but only two are found
in Herrerasaurus);[7] and a perforate acetabulum, or hip socket, with a hole at the center of
its inside surface (closed in Saturnalia tupiniquim, for example).[30][31] Another difficulty of
determining distinctly dinosaurian features is that early dinosaurs and other archosaurs
from the Late Triassic epoch are often poorly known and were similar in many ways; these
animals have sometimes been misidentified in the literature. [32]
Dinosaurs stand with their hind limbs erect in a manner similar to most modern mammals,
but distinct from most other reptiles, whose limbs sprawl out to either side.[33] This posture is
due to the development of a laterally facing recess in the pelvis (usually an open socket)
and a corresponding inwardly facing distinct head on the femur.[34] Their erect posture
enabled early dinosaurs to breathe easily while moving, which likely permitted stamina and
activity levels that surpassed those of "sprawling" reptiles.[35] Erect limbs probably also
helped support the evolution of large size by reducing bending stresses on limbs. [36] Some
non-dinosaurian archosaurs, including rauisuchians, also had erect limbs but achieved this
by a "pillar-erect" configuration of the hip joint, where instead of having a projection from
the femur insert on a socket on the hip, the upper pelvic bone was rotated to form an
overhanging shelf.[36]

History of study
Further information: History of paleontology

Pre-scientific history
Dinosaur fossils have been known for millennia, although their true nature was not
recognized. The Chinese considered them to be dragon bones and documented them as
such. For example, Huayang Guo Zhi (華陽國志), a gazetteer compiled by Chang Qu (常璩
) during the Western Jin Dynasty (265–316), reported the discovery of dragon bones at
Wucheng in Sichuan Province.[37] Villagers in central China have long unearthed fossilized
"dragon bones" for use in traditional medicines.[38] In Europe, dinosaur fossils were
generally believed to be the remains of giants and other biblical creatures.[39]

Early dinosaur research

William Buckland

Scholarly descriptions of what would now be recognized as dinosaur bones first appeared
in the late 17th century in England. Part of a bone, now known to have been the femur of
a Megalosaurus,[40] was recovered from a limestone quarry at Cornwell near Chipping
Norton, Oxfordshire, in 1676. The fragment was sent to Robert Plot, Professor of Chemistry
at the University of Oxford and first curator of the Ashmolean Museum, who published a
description in his The Natural History of Oxford-shire (1677).[41] He correctly identified the
bone as the lower extremity of the femur of a large animal, and recognized that it was too
large to belong to any known species. He, therefore, concluded it to be the femur of a huge
human, perhaps a Titan or another type of giant featured in legends.[42][43] Edward Lhuyd, a
friend of Sir Isaac Newton, published Lithophylacii Britannici ichnographia (1699), the first
scientific treatment of what would now be recognized as a dinosaur when he described and
named a sauropod tooth, "Rutellum impicatum",[44][45] that had been found in Caswell,
near Witney, Oxfordshire.[46]
Sir Richard Owen's coining of the word dinosaur, at a meeting of the British Association for the
Advancement of Science in 1841

Between 1815 and 1824, the Rev William Buckland, the first Reader of Geology at the
University of Oxford, collected more fossilized bones of Megalosaurus and became the first
person to describe a non-avian dinosaur in a scientific journal.[40][47] The second non-avian
dinosaur genus to be identified, Iguanodon, was discovered in 1822 by Mary Ann Mantell –
the wife of English geologist Gideon Mantell. Gideon Mantell recognized similarities
between his fossils and the bones of modern iguanas. He published his findings in
1825.[48][49]
The study of these "great fossil lizards" soon became of great interest to European and
American scientists, and in 1841 the English paleontologist Sir Richard Owen coined the
term "dinosaur", using it to refer to the "distinct tribe or sub-order of Saurian Reptiles" that
were then being recognized in England and around the world.[50][51] The term is derived
from Ancient Greek δεινός (deinos) 'terrible, potent or fearfully great',
and σαῦρος (sauros) 'lizard or reptile'.[50][52] Though the taxonomic name has often been
interpreted as a reference to dinosaurs' teeth, claws, and other fearsome characteristics,
Owen intended it to also evoke their size and majesty. [53] Owen recognized that the remains
that had been found so far, Iguanodon, Megalosaurus and Hylaeosaurus, shared a number
of distinctive features, and so decided to present them as a distinct taxonomic group. With
the backing of Prince Albert, the husband of Queen Victoria, Owen established the Natural
History Museum, London, to display the national collection of dinosaur fossils and other
biological and geological exhibits.[54]

Discoveries in North America

In 1858, William Parker Foulke discovered the first known American dinosaur, in marl pits
in the small town of Haddonfield, New Jersey. (Although fossils had been found before,
their nature had not been correctly discerned.) The creature was named Hadrosaurus
foulkii. It was an extremely important find: Hadrosaurus was one of the first nearly complete
dinosaur skeletons found (the first was in 1834, in Maidstone, England), and it was clearly a
bipedal creature. This was a revolutionary discovery as, until that point, most scientists had
believed dinosaurs walked on four feet, like other lizards. Foulke's discoveries sparked a
wave of interests in dinosaurs in the United States, known as dinosaur mania.[55]
 Edward Drinker Cope

Othniel Charles Marsh

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

"Dinosaur renaissance" and beyond

Main article: Dinosaur renaissance

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 and 1969 description
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.[57][58][59][60][61][62] 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 well-preserved feathered dinosaurs 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.[63]

Soft tissue and DNA

Scipionyx fossil with intestines, Natural History Museum of Milan

One of the best examples of soft-tissue impressions in a fossil dinosaur was discovered in
the Pietraroia Plattenkalk in southern Italy. The discovery was reported in 1998, and
described the specimen of a small, juvenile coelurosaur, Scipionyx samniticus. The fossil
includes portions of the intestines, colon, liver, muscles, and windpipe of this dinosaur. [64]
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. [65] 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.[65]
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 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. [66]
The extraction of ancient DNA from dinosaur fossils has been reported on two separate
occasions;[67] upon further inspection and peer review, however, neither of these reports
could be confirmed.[68] 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. [69] In addition,
several proteins, including hemoglobin,[70] have putatively been detected in dinosaur
fossils.[71][72]
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.[73][74]

Evolutionary history
Origins and early evolution
The early dinosaurs Herrerasaurus (large), Eoraptor (small) and a Plateosaurus skull, from
the Triassic

Dinosaurs diverged from their archosaur ancestors during the Middle to Late Triassic
epochs, roughly 20 million years after the devastating Permian–Triassic extinction
event wiped out an estimated 96% of all marine species and 70% of terrestrial vertebrate
species approximately 252 million years ago.[75][76] Radiometric dating of the Ischigualasto
Formation of Argentina where the early dinosaur genus Eoraptor was found date it as
231.4 million years old.[77] Eoraptor is thought to resemble the common ancestor of all
dinosaurs; if this is true, its traits suggest that the first dinosaurs were small,
bipedal predators.[78][79][80] The discovery of primitive, dinosaur-like ornithodirans such
as Lagosuchus and Lagerpeton in Argentina in the Carnian epoch of the Triassic, around
233 million years ago,[81] supports this view; analysis of recovered fossils suggests that
these animals were indeed small, bipedal predators. Dinosaurs may have appeared as
early as the Anisian epoch of the Triassic, 245 million years ago, as evidenced by remains
of the genus Nyasasaurus from that period. However, its known fossils are too fragmentary
to tell if it was a dinosaur or only a close relative. [82] Paleontologist Max C. Langer et
al. (2018) determined that Staurikosaurus from the Santa Maria Formation dates to
233.23 million years ago, making it older in geologic age than Eoraptor.[83]
When dinosaurs appeared, they were not the dominant terrestrial animals. The terrestrial
habitats were occupied by various types of archosauromorphs and therapsids,
like cynodonts and rhynchosaurs. Their main competitors were the pseudosuchians, such
as aetosaurs, ornithosuchids and rauisuchians, which were more successful than the
dinosaurs.[84] Most of these other animals became extinct in the Triassic, in one of two
events. First, at about 215 million years ago, a variety of basal archosauromorphs,
including the protorosaurs, became extinct. This was followed by the Triassic–Jurassic
extinction event (about 201 million years ago), that saw the end of most of the other groups
of early archosaurs, like aetosaurs, ornithosuchids, phytosaurs, and rauisuchians.
Rhynchosaurs and dicynodonts survived (at least in some areas) at least as late as early –
mid Norian and late Norian or earliest Rhaetian stages, respectively,[85][86] and the exact date
of their extinction is uncertain. These losses left behind a land fauna of crocodylomorphs,
dinosaurs, mammals, pterosaurians, and turtles.[7] The first few lines of early
dinosaurs diversified through the Carnian and Norian stages of the Triassic, possibly by
occupying the niches of the groups that became extinct. [9] Also notably, there was a
heightened rate of extinction during the Carnian pluvial event.[87]

Evolution and paleobiogeography

The supercontinent Pangaea in the early Mesozoic (around 200 million years ago)

Dinosaur evolution after the Triassic followed changes in vegetation and the location of
continents. In the Late Triassic and Early Jurassic, the continents were connected as the
single landmass Pangaea, and there was a worldwide dinosaur fauna mostly composed
of coelophysoid carnivores and early sauropodomorph herbivores. [88] 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.[89] The general homogeneity of dinosaurian faunas continued into the Middle
and Late Jurassic, where most localities had predators consisting
of ceratosaurians, megalosauroids, and allosauroids, and herbivores consisting of
stegosaurian ornithischians and large sauropods. Examples of this include the Morrison
Formation of North America and Tendaguru Beds of Tanzania. Dinosaurs in China show
some differences, with specialized metriacanthosaurid theropods and unusual, long-necked
sauropods like Mamenchisaurus.[88] Ankylosaurians and ornithopods were also becoming
more common, but primitive sauropodomorphs had become extinct. Conifers
and pteridophytes were the most common plants. Sauropods, like earlier
sauropodomorphs, were not oral processors, but ornithischians were evolving various
means of dealing with food in the mouth, including potential cheek-like organs to keep food
in the mouth, and jaw motions to grind food. [89] Another notable evolutionary event of the
Jurassic was the appearance of true birds, descended from maniraptoran
coelurosaurians.[11]
By the Early Cretaceous and the ongoing breakup of Pangaea, dinosaurs were becoming
strongly differentiated by landmass. The earliest part of this time saw the spread of
ankylosaurians, iguanodontians, and brachiosaurids through Europe, North America, and
northern Africa. These were later supplemented or replaced in Africa by large spinosaurid
and carcharodontosaurid theropods, and rebbachisaurid and titanosaurian sauropods, also
found in South America. In Asia, maniraptoran coelurosaurians
like dromaeosaurids, troodontids, and oviraptorosaurians became the common theropods,
and ankylosaurids and early ceratopsians like Psittacosaurus became important
herbivores. Meanwhile, Australia was home to a fauna of basal
ankylosaurians, hypsilophodonts, and iguanodontians.[88] The stegosaurians appear to have
gone extinct at some point in the late Early Cretaceous or early Late Cretaceous. A major
change in the Early Cretaceous, which would be amplified in the Late Cretaceous, was the
evolution of flowering plants. At the same time, several groups of dinosaurian herbivores
evolved more sophisticated ways to orally process food. Ceratopsians developed a method
of slicing with teeth stacked on each other in batteries, and iguanodontians refined a
method of grinding with dental batteries, taken to its extreme in hadrosaurids.[89] Some
sauropods also evolved tooth batteries, best exemplified by the
rebbachisaurid Nigersaurus.[90]
There were three general dinosaur faunas in the Late Cretaceous. In the northern
continents of North America and Asia, the major theropods were tyrannosaurids and
various types of smaller maniraptoran theropods, with a predominantly ornithischian
herbivore assemblage of hadrosaurids, ceratopsians, ankylosaurids, and
pachycephalosaurians. In the southern continents that had made up the now-splitting
supercontinent Gondwana, abelisaurids were the common theropods, and titanosaurian
sauropods the common herbivores. Finally, in Europe,
dromaeosaurids, rhabdodontid iguanodontians, nodosaurid ankylosaurians, and
titanosaurian sauropods were prevalent.[88] Flowering plants were greatly radiating,[89] with
the first grasses appearing by the end of the Cretaceous.[91] Grinding hadrosaurids and
shearing ceratopsians became very diverse across North America and Asia. Theropods
were also radiating as herbivores or omnivores,
with therizinosaurians and ornithomimosaurians becoming common.[89]
The Cretaceous–Paleogene extinction event, which occurred approximately 66 million
years ago at the end of the Cretaceous, caused the extinction of all dinosaur groups except
for the neornithine birds. Some other diapsid groups,
including crocodilians, dyrosaurs, sebecosuchians,
turtles, lizards, snakes, sphenodontians, and choristoderans, also survived the event.[92]
The surviving lineages of neornithine birds, including the ancestors of modern ratites, ducks
and chickens, and a variety of waterbirds, diversified rapidly at the beginning of
the Paleogene period, entering ecological niches left vacant by the extinction of Mesozoic
dinosaur groups such as the arboreal enantiornithines, aquatic hesperornithines, and even
the larger terrestrial theropods (in the form of Gastornis, eogruiids, bathornithids,
ratites, geranoidids, mihirungs, and "terror birds"). It is often stated that mammals out-
competed the neornithines for dominance of most terrestrial niches but many of these
groups co-existed with rich mammalian faunas for most of the Cenozoic Era.[93] Terror birds
and bathornithids occupied carnivorous guilds alongside predatory mammals, [94][95] and
ratites are still fairly successful as mid-sized herbivores; eogruiids similarly lasted from
the Eocene to Pliocene, only becoming extinct very recently after over 20 million years of
co-existence with many mammal groups.[96]

Classification
Main article: Dinosaur classification

Saurischian pelvis structure (left side)

Tyrannosaurus pelvis (showing saurischian structure – left side)

Ornithischian pelvis structure (left side)

Edmontosaurus pelvis (showing ornithischian structure – left side)

Dinosaurs belong to a group known as archosaurs, which also includes modern

crocodilians. Within the archosaur group, dinosaurs are differentiated most noticeably by
their gait. Dinosaur legs extend directly beneath the body, whereas the legs of lizards and
crocodilians sprawl out to either side.[27]
Collectively, dinosaurs as a clade are divided into two primary branches, Saurischia and
Ornithischia. Saurischia includes those taxa sharing a more recent common ancestor with
birds than with Ornithischia, while Ornithischia includes all taxa sharing a more recent
common ancestor with Triceratops than with Saurischia. Anatomically, these two groups
can be distinguished most noticeably by their pelvic structure. Early saurischians—"lizard-
hipped", from the Greek sauros (σαῦρος) meaning "lizard" and ischion (ἰσχίον) meaning
"hip joint"—retained the hip structure of their ancestors, with a pubis bone
directed cranially, or forward.[34] This basic form was modified by rotating the pubis
backward to varying degrees in several groups
(Herrerasaurus,[97] therizinosauroids,[98] dromaeosaurids,[99] and birds[11]). Saurischia includes
the theropods (exclusively bipedal and with a wide variety of diets) and sauropodomorphs
(long-necked herbivores which include advanced, quadrupedal groups). [26][100]
By contrast, ornithischians—"bird-hipped", from the Greek ornitheios (ὀρνίθειος) meaning
"of a bird" and ischion (ἰσχίον) meaning "hip joint"—had a pelvis that superficially
resembled a bird's pelvis: the pubic bone was oriented caudally (rear-pointing). Unlike
birds, the ornithischian pubis also usually had an additional forward-pointing process.
Ornithischia includes a variety of species that were primarily herbivores.
Despite the terms "bird hip" (Ornithischia) and "lizard hip" (Saurischia), birds are not part of
Ornithischia. Birds instead belong to Saurischia, the “lizard-hipped” dinosaurs—birds
evolved from earlier dinosaurs with "lizard hips". [27]

Taxonomy
The following is a simplified classification of dinosaur groups based on their evolutionary
relationships, and those of the main dinosaur groups Theropoda, Sauropodomorpha and
Ornithischia, exemplified by the studies of Molina-Pérez and Larramendi in 2019 for
Theropoda and 2020 for Sauropodomorpha,[101][102] and Madzia and colleagues in 2021 for
Ornithischia.[103] The dagger (†) is used to signify groups with no living members,
descriptions of clades follow Holtz (2007).[104][105] The dagger (†) is used to signify groups
with no living members.

 Dinosauria

 Saurischia

 †Herrerasauria (early bipedal carnivores)[106]

 Theropoda (carnivorous)

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

 †Dilophosauridae (larger kink-snouted dinosaurs, previously considered
coelophysoids)
 †Ceratosauria (generally elaborately horned, power carnivores that existed
from the Jurassic to Cretaceous periods, originally included Coelophysoidea)

 †Ceratosauridae (A group of Ceratosaurs, they existed in North and South

America and possibly even Africa during the Jurassic and Early Cretaceous)
 †Abelisauroidea (A group of Ceratosaurs that include the Abelisauridae and the
Noasauridae, they were the dominant Carnivores in the Southeren hemisphere
at the end of the Cretaceous)[107]

 Tetanurae (stiff-tailed dinosaurs)

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

spinosaurids, possibly basal members of Carnosauria) [108]
 †Carnosauria (giant meat-eating dinosaurs; Allosaurus and close relatives,
like Carcharodontosaurus)
 Coelurosauria (feathered theropods, with a range of body sizes and niches) [63]

 †?Megaraptora (theropods with large hand claws)

 †Compsognathidae (small early coelurosaurs with short forelimbs)
 †Tyrannosauroidea (Tyrannosaurus and close relatives)
 †Ornithomimosauria (ostrich dinosaurs; small-headed, mostly toothless,
omnivorous or possible herbivores)
 Maniraptora (feathered dinosaurs)

Restoration of six dromaeosaurid theropods: from left to

right Microraptor, Velociraptor, Austroraptor, Dromaeosaurus, Utahraptor, and Deinonychus

 †Alvarezsauroidea (small insectivores with reduced forelimbs each bearing one

enlarged claw)
 †Therizinosauria (sloth dinosaurs)
 †Oviraptorosauria (egg-thief dinosaurs)
 †Dromaeosauridae (raptor dinosaurs; heavier, shorter legs and longer arms)
 †Troodontidae (long-legged raptor dinosaurs)
 †Scansoriopterygidae (small primitive avialans with long third fingers)
 Avialae (modern birds and extinct relatives)

 †Archaeopterygidae (small, winged theropods or primitive birds)

 †Confuciusornithidae (small toothless avialans)
 †Enantiornithes (primitive tree-dwelling, flying avialans)
 Pygostylia (advanced flying birds)

 †Yanornithidae
 †Songlingornithidae
 †Hesperornithes (specialized aquatic diving birds)
 Neornithes (modern, beaked birds and their extinct relatives)
 Restoration of four macronarian sauropods: from left to
right Camarasaurus, Brachiosaurus, Giraffatitan, and Euhelopus

 †Sauropodomorpha (herbivores with small heads, long necks, long tails)

 †Plateosauridae (primitive, strictly bipedal "prosauropods")

 †Riojasauridae (large, primitive sauropodomorphs)
 †Massospondylidae (long-necked, primitive sauropodomorphs)
 †Sauropoda (very large and heavy; quadrupedal)

 †Mamenchisauridae (group of Sauropods with very long, thin necks)

 †Cetiosauridae ("whale reptiles")
 †Turiasauria (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)

 †Euhelopodidae (bizarre stocky Macronarians)[109]
 †Titanosauria (diverse; stocky, with wide hips; most common in the Late
Cretaceous of southern continents)

 †Aeolosaurini
 †Saltasauroidea

 †Saltasaurinae
 †Lognkosauria

Restoration of six ornithopods; far left: Camptosaurus, left: Iguanodon, center

background: Shantungosaurus, center foreground: Dryosaurus,
right: Corythosaurus, far right (large) Tenontosaurus.

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

 †Heterodontosauridae (small basal ornithopod herbivores/omnivores with

prominent canine-like teeth)
 †Thyreophora (armored dinosaurs; mostly quadrupeds)
 †Ankylosauria (scutes as primary armor; some had club-like tails)
 †Stegosauria (spikes and plates as primary armor)

 †Neornithischia ("new ornithischians")

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

chewing using skull flexibility and numerous teeth)

 †Elasmaria
 †Iguanodontia (contains most Ornithopod groups)

 †Rhabdodontomorpha (A group of Ornithopods which existed from the Early to

Late Cretaceous period)[110]
 †Dryosauridae (A group of small Ornithopods from the Jurassic to Late
Cretacoues)
 †Ankylopollexia (A group that contains most of the Large Ornithopods,
ancestrally had a thumb spike)[111]

 †Camptosauridae
 †Hadrosauroidea (large quadrupedal herbivores, with teeth merged into dental
batteries)

 †Marginocephalia (characterized by a cranial growth)

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

 †Ceratopsia (bipeds and quadrupeds with neck frills; many also had horns)
Timeline of major groups
Timeline of major dinosaur groups per Holtz (2007).
Paleobiology
Knowledge about dinosaurs is derived from a variety of fossil and non-
fossil records, including fossilized
bones, feces, trackways, gastroliths, feathers, impressions of skin, internal
organs and other soft tissues.[64][65] 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).[112][113] Two
topics of particular interest and study have been dinosaur size and
behavior.[114]

Size
Main article: Dinosaur size
 Scale diagram comparing the average human to the longest known dinosaurs
in five major clades:
Sauropoda (Supersaurus vivianae)
Ornithopoda (Shantungosaurus giganteus)
Theropoda (Spinosaurus aegyptiacus)
Thyreophora (Stegosaurus ungulatus)
Marginocephalia (Triceratops prorsus)

Current evidence suggests that dinosaur average size varied through the
Triassic, Early Jurassic, Late Jurassic and Cretaceous. [79] 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.[115] The mode of
Mesozoic dinosaur body masses is between 1 to 10 metric tons (1.1 to
11.0 short tons).[116] This contrasts sharply with the average size of
Cenozoic mammals, estimated by the National Museum of Natural
History as about 2 to 5 kg (4.4 to 11.0 lb).[117]
The sauropods were the largest and heaviest dinosaurs. For much of the
dinosaur era, the smallest sauropods were larger than anything else in
their habitat, and the largest was an order of magnitude more massive
than anything else that has since walked the Earth. Giant prehistoric
mammals such as Paraceratherium (the largest land mammal ever) were
dwarfed by the giant sauropods, and only modern whales approach or
surpass them in size.[118] 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. [119]
Largest and smallest
Scientists will probably never be certain of the largest and smallest
dinosaurs to have ever existed. This is because only a tiny percentage of
animals were ever fossilized and most of these remain buried in the earth.
Few of the specimens that are recovered are complete skeletons, and
impressions of skin and other soft tissues are rare. Rebuilding a complete
skeleton by comparing the size and morphology of bones to those of
similar, better-known species is an inexact art, and reconstructing the
muscles and other organs of the living animal is, at best, a process of
educated guesswork.[120]
Comparative size of Argentinosaurus to the average human

The tallest and heaviest dinosaur known from good skeletons

is Giraffatitan brancai (previously classified as a species
of Brachiosaurus). Its remains were discovered in Tanzania between 1907
and 1912. Bones from several similar-sized individuals were incorporated
into the skeleton now mounted and on display at the Museum für
Naturkunde in Berlin;[121] this mount is 12 meters (39 ft) tall and 21.8 to 22.5
meters (72 to 74 ft) long,[122][123] and would have belonged to an animal that
weighed between 30000 and 60000 kilograms (70000 and 130000 lb). The
longest complete dinosaur is the 27 meters (89 ft) long Diplodocus, which
was discovered in Wyoming in the United States and displayed
in Pittsburgh's Carnegie Museum of Natural History in 1907.[124] The
longest dinosaur known from good fossil material is the Patagotitan: the
skeleton mount in the American Museum of Natural History in New York is
37 meters (121 ft) long. The Museo Municipal Carmen Funes in Plaza
Huincul, Argentina, has an Argentinosaurus reconstructed skeleton mount
that is 39.7 meters (130 ft) long.[125]

An adult bee hummingbird, the smallest known dinosaur

There were larger dinosaurs, but knowledge of them is based entirely on a

small number of fragmentary fossils. Most of the largest herbivorous
specimens on record were discovered in the 1970s or later, and include
the massive Argentinosaurus, which may have
weighed 80000 to 100000 kilograms (90 to 110 short tons) and reached
lengths of 30 to 40 meters (98 to 131 ft); some of the longest were the
33.5-meter (110 ft) long Diplodocus hallorum[119] (formerly Seismosaurus),
the 33-to-34-meter (108 to 112 ft) long Supersaurus,[126] and 37-meter
(121 ft) long Patagotitan; and the tallest, the 18-meter (59 ft)
tall Sauroposeidon, which could have reached a sixth-floor window. The
heaviest and longest dinosaur may have been Maraapunisaurus, known
only from a now lost partial vertebral neural arch described in 1878.
Extrapolating from the illustration of this bone, the animal may have been
58 meters (190 ft) long and weighed 122400 kg (270000 lb).[119] However,
as no further evidence of sauropods of this size has been found, and the
discoverer, Cope, had made typographic errors before, it is likely to have
been an extreme overestimation.[127]
The largest carnivorous dinosaur was Spinosaurus, reaching a length of
12.6 to 18 meters (41 to 59 ft), and weighing 7 to 20.9 metric tons (7.7 to
23.0 short tons).[128][129] Other large carnivorous theropods
included Giganotosaurus, Carcharodontosaurus and Tyrannosaurus.[129] Th
erizinosaurus and Deinocheirus were among the tallest of the theropods.
The largest ornithischian dinosaur was probably the
hadrosaurid Shantungosaurus giganteus which measured 16.6 meters
(54 ft).[130] The largest individuals may have weighed as much as 16 metric
tons (18 short tons).[131]
The smallest dinosaur known is the bee hummingbird,[132] with a length of
only 5 centimeters (2.0 in) and mass of around 1.8 g (0.063 oz).[133] The
smallest known non-avialan dinosaurs were about the size of pigeons and
were those theropods most closely related to birds. [134] For
example, Anchiornis huxleyi is currently the smallest non-avialan dinosaur
described from an adult specimen, with an estimated weight of 110 g
(3.9 oz)[135] and a total skeletal length of 34 centimeters (1.12 ft).[134][135] The
smallest herbivorous non-avialan dinosaurs
included Microceratus and Wannanosaurus, at about 60 centimeters
(2.0 ft) long each.[104][136]

Behavior

A nesting ground of the hadrosaur Maiasaura peeblesorum was discovered in

1978

Many modern birds are highly social, often found living in flocks. There is
general agreement that some behaviors that are common in birds, as well
as in crocodiles (closest living relatives of birds), were also common
among extinct dinosaur groups. Interpretations of behavior in fossil species
are generally based on the pose of skeletons and their habitat, computer
simulations of their biomechanics, and comparisons with modern animals
in similar ecological niches.[112]
The first potential evidence for herding or flocking as a widespread
behavior common to many dinosaur groups in addition to birds was the
1878 discovery of 31 Iguanodon, ornithischians that were then thought to
have perished together in Bernissart, Belgium, after they fell into a deep,
flooded sinkhole and drowned.[137] Other mass-death sites have been
discovered subsequently. Those, along with multiple trackways, suggest
that gregarious behavior was common in many early dinosaur species.
Trackways of hundreds or even thousands of herbivores indicate that
duck-billed (hadrosaurids) may have moved in great herds, like
the American bison or the African Springbok. Sauropod tracks document
that these animals traveled in groups composed of several different
species, at least in Oxfordshire, England,[138] although there is no evidence
for specific herd structures.[139] Congregating into herds may have evolved
for defense, for migratory purposes, or to provide protection for young.
There is evidence that many types of slow-growing dinosaurs, including
various theropods, sauropods, ankylosaurians, ornithopods, and
ceratopsians, formed aggregations of immature individuals. One example
is a site in Inner Mongolia that has yielded remains of over
20 Sinornithomimus, from one to seven years old. This assemblage is
interpreted as a social group that was trapped in mud. [140] The interpretation
of dinosaurs as gregarious has also extended to depicting carnivorous
theropods as pack hunters working together to bring down large
prey.[141][142] 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.[143]

Restoration of two Centrosaurus apertus engaged in intra-specific combat

The crests and frills of some dinosaurs, like the marginocephalians,

theropods and lambeosaurines, may have been too fragile to be used for
active defense, and so they were likely used for sexual or aggressive
displays, though little is known about dinosaur mating and territorialism.
Head wounds from bites suggest that theropods, at least, engaged in
active aggressive confrontations. [144]
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,[145] providing evidence that
dinosaurs did indeed attack each other. [146] 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. [146] Cannibalism amongst
some species of dinosaurs was confirmed by tooth marks found
in Madagascar in 2003, involving the theropod Majungasaurus.[147]
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.[148]
Based on fossil evidence from dinosaurs such as Oryctodromeus, some
ornithischian species seem to have led a partially fossorial (burrowing)
lifestyle.[149] Many modern birds are arboreal (tree climbing), and this was
also true of many Mesozoic birds, especially the enantiornithines. [150] While
some early bird-like species may have already been arboreal as well
(including dromaeosaurids) such as Microraptor[151]) 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,[112] whether diplodocids could create sonic booms via whip-like tail
snapping,[152] and whether sauropods could float.[153]

Communication

Restoration of a striking and unusual visual display in a Lambeosaurus

magnicristatus

Modern birds are known to communicate using visual and auditory signals,
and the wide diversity of visual display structures among fossil dinosaur
groups, such as horns, frills, crests, sails, and feathers, suggests that
visual communication has always been important in dinosaur
biology.[154] Reconstruction of the plumage color of Anchiornis, suggest the
importance of color in visual communication in non-avian
dinosaurs.[155] The evolution of dinosaur vocalization is less certain.
Paleontologist Phil Senter has suggested that non-avian dinosaurs relied
mostly on visual displays and possibly non-vocal acoustic sounds like
hissing, jaw grinding or clapping, splashing and wing beating (possible in
winged maniraptoran dinosaurs). He states they were unlikely to have
been capable of vocalizing since their closest relatives, crocodilians and
birds, use different means to vocalize, the former via the larynx and the
latter through the unique syrinx, suggesting they evolved independently
and their common ancestor was mute. [154]
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 years ago, and this organ is unlikely to have existed in non-
avian dinosaurs. However, in contrast to Senter, other 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.[156] A 2016 study suggests that some
dinosaurs produced closed mouth vocalizations like cooing, hooting and
booming. These occur in both reptiles and birds and involve inflating the
esophagus or tracheal pouches. Such vocalizations evolved independently
in extant archosaurs numerous times, following increases in body
size.[157] The crests of the Lambeosaurini and nasal chambers of
ankylosaurids have been suggested to have functioned in vocal
resonance,[158][159] though Senter stated 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. [154]

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

All dinosaurs laid amniotic eggs. Dinosaur eggs were usually laid in a nest.
Most species create somewhat elaborate nests which can be cups,
domes, plates, beds scrapes, mounds, or burrows. [160] 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.[161]
When laying eggs, females grow a special type of bone between the hard
outer bone and the marrow of their limbs. This medullary bone, which is
rich in calcium, is used to make eggshells. A discovery of features in
a Tyrannosaurus skeleton provided evidence of medullary bone in extinct
dinosaurs and, for the first time, allowed paleontologists to establish the
sex of a fossil dinosaur specimen. Further research has found medullary
bone in the carnosaur Allosaurus and the ornithopod Tenontosaurus.
Because the line of dinosaurs that
includes Allosaurus and Tyrannosaurus diverged from the line that led
to Tenontosaurus very early in the evolution of dinosaurs, this suggests
that the production of medullary tissue is a general characteristic of all
dinosaurs.[162]

Fossil interpreted as a nesting oviraptorid Citipati at the American Museum of

Natural History. Smaller fossil far right showing inside one of the eggs.

Another widespread trait among modern birds (but see below in regards to
fossil groups and extant megapodes) is parental care for young after
hatching. Jack Horner's 1978 discovery of a Maiasaura ("good mother
lizard") nesting ground in Montana demonstrated that parental care
continued long after birth among ornithopods.[163] A specimen of
the oviraptorid Citipati osmolskae was discovered in a chicken-like
brooding position in 1993,[164] which may indicate that they had begun using
an insulating layer of feathers to keep the eggs warm. [165] An embryo of the
basal sauropodomorph Massospondylus was found without teeth,
indicating that some parental care was required to feed the young
dinosaurs.[166] Trackways have also confirmed parental behavior among
ornithopods from the Isle of Skye in northwestern Scotland.[167]
However, there is ample evidence
of precociality or superprecociality among many dinosaur species,
particularly theropods. For instance, non-ornithuromorph birds have been
abundantly demonstrated to have had slow growth rates, megapode-like
egg burying behavior and the ability to fly soon after
birth.[168][169][170][171] Both Tyrannosaurus and Troodon had juveniles with clear
superprecociality and likely occupying different ecological niches than the
adults.[161] Superprecociality has been inferred for sauropods. [172]
Genital structures are unlikely to fossilize as they lack scales that may
allow preservation via pigmentation or residual calcium phosphate salts. In
2021, the best preserved specimen of a dinosaur's cloacal vent exterior
was described for Psittacosaurus, demonstrating lateral swellings similar to
crocodylian musk glands used in social displays by both sexes and
pigmented regions which could also reflect a signalling function. However,
this specimen on its own does not offer enough information to determine
whether this dinosaur had sexual signalling functions; it only supports the
possibility. Cloacal visual signalling can occur in either males or females in
living birds, making it unlikely to be useful to determine sex for extinct
dinosaurs.[173]

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. [174] While all modern birds have high
metabolisms and are endothermic ("warm-blooded"), a vigorous debate
has been ongoing since the 1960s regarding how far back in the dinosaur
lineage this trait extended. Various researchers have supported dinosaurs
as being endothermic, ectothermic ("cold-blooded"), or somewhere in
between.[175] An emerging consensus among researchers is that, while
different lineages of dinosaurs would have had different metabolisms, most
of them had higher metabolic rates than other reptiles but lower than living
birds and mammals,[176] which is termed mesothermy by some.[177] Evidence
from crocodiles and their extinct relatives suggests that such elevated
metabolisms could have developed in the earliest archosaurs, which were
the common ancestors of dinosaurs and crocodiles. [178][179]

This 1897 restoration of Brontosaurus as an aquatic, tail-dragging animal,

by Charles R. Knight, typified early views on dinosaur lifestyles.

After non-avian dinosaurs were discovered, paleontologists first posited

that they were ectothermic. This was used to imply that the ancient
dinosaurs were relatively slow, sluggish organisms, even though many
modern reptiles are fast and light-footed despite relying on external
sources of heat to regulate their body temperature. The idea of dinosaurs
as ectothermic remained a prevalent view until Robert T. Bakker, an early
proponent of dinosaur endothermy, published an influential paper on the
topic in 1968. Bakker specifically used anatomical and ecological evidence
to argue that sauropods, which had hitherto been depicted as sprawling
aquatic animals with their tails dragging on the ground, were endotherms
that lived vigorous, terrestrial lives. In 1972, Bakker expanded on his
arguments based on energy requirements and predator-prey ratios. This
was one of the seminal results that led to the Dinosaur
renaissance.[58][59][61][180]
One of the greatest contributions to the modern understanding of dinosaur
physiology has been paleohistology, the study of microscopic tissue
structure in dinosaurs.[181][182] From the 1960s forward, Armand de
Ricqlès suggested that the presence of fibrolamellar bone—bony tissue
with an irregular, fibrous texture and filled with blood vessels—was
indicative of consistently fast growth and therefore endothermy.
Fibrolamellar bone was common in both dinosaurs and
pterosaurs,[183][184] though not universally present.[185][186] This has led to a
significant body of work in reconstructing growth curves and modeling the
evolution of growth rates across various dinosaur lineages, [187] which has
suggested overall that dinosaurs grew faster than living reptiles. [182] Other
lines of evidence suggesting endothermy include the presence of feathers
and other types of body coverings in many lineages (see § Feathers);
more consistent ratios of the isotope oxygen-18 in bony tissue compared
to ectotherms, particularly as latitude and thus air temperature varied,
which suggests stable internal temperatures [188][189] (although these ratios
can be altered during fossilization[190]); and the discovery of polar
dinosaurs, which lived in Australia, Antarctica, and Alaska when these
places would have had cool, temperate climates.[191][192][193][194]

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. [195] Such respiratory systems, which
may have appeared in the earliest saurischians, [196] would have provided
them with more oxygen compared to a mammal of similar size, while also
having a larger resting tidal volume and requiring a lower breathing
frequency, which would have allowed them to sustain higher activity
levels.[118] The rapid airflow would also have been an effective cooling
mechanism, which in conjunction with a lower metabolic rate[197] would
have prevented large sauropods from overheating. These traits may have
enabled sauropods to grow quickly to gigantic sizes. [198][199] Sauropods may
also have benefitted from their size—their small surface area to volume
ratio meant that they would have been able to thermoregulate more easily,
a phenomenon termed gigantothermy.[118][200]
Like other reptiles, dinosaurs are primarily uricotelic, that is,
their kidneys extract nitrogenous wastes from their bloodstream and
excrete it as uric acid instead of urea or ammonia via the ureters into the
intestine. This would have helped them to conserve water. [176] In most living
species, uric acid is excreted along with feces as a semisolid
waste.[201][202] However, at least some modern birds (such as hummingbirds)
can be facultatively ammonotelic, excreting most of the nitrogenous
wastes as ammonia.[203] This material, as well as the output of the
intestines, emerges from the cloaca.[204][205] In addition, many species
regurgitate pellets,[206] and fossil pellets are known as early as the Jurassic
from Anchiornis.[207]
The size and shape of the brain can be partly reconstructed based on the
surrounding bones. In 1896, Marsh calculated ratios between brain weight
and body weight of seven species of dinosaurs, showing that the brain of
dinosaurs was proportionally smaller than in today's crocodiles, and that
the brain of Stegosaurus was smaller than in any living land vertebrate.
This contributed to the widespread public notion of dinosaurs as being
sluggish and extraordinarily stupid. Harry Jerison, in 1973, showed that
proportionally smaller brains are expected at larger body sizes, and that
brain size in dinosaurs was not smaller than expected when compared to
living reptiles.[208] Later research showed that relative brain size
progressively increased during the evolution of theropods, with the highest
intelligence – comparable to that of modern birds – calculated for the
troodontid Troodon.[209]

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.[210] After the work of Gerhard
Heilmann in the early 20th century, the theory of birds as dinosaur
descendants was abandoned in favor of the idea of them being
descendants of generalized thecodonts, with the key piece of evidence
being the supposed lack of clavicles in dinosaurs.[211] However, as later
discoveries showed, clavicles (or a single fused wishbone, which derived
from separate clavicles) were not actually absent; [11] they had been found
as early as 1924 in Oviraptor, but misidentified as an interclavicle.[212] In the
1970s, Ostrom revived the dinosaur–bird theory,[213] which gained
momentum in the coming decades with the advent of cladistic
analysis,[214] and a great increase in the discovery of small theropods and
early birds.[29] 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.[63][11] Birds share over a hundred distinct
anatomical features with theropod dinosaurs, which are now generally
accepted to have been their closest ancient relatives.[215] They are most
closely allied with maniraptoran coelurosaurs.[11] 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,[216] or that maniraptoran theropods are the ancestors of
birds but themselves are not dinosaurs, only convergent with dinosaurs.[217]

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 also shared by several non-avian dinosaurs. Based on
the current distribution of fossil evidence, it appears that feathers were an
ancestral dinosaurian trait, though one that may have been selectively lost
in some species.[218] Direct fossil evidence of feathers or feather-like
structures has been discovered in a diverse array of species in many non-
avian dinosaur groups,[63] both among saurischians and ornithischians.
Simple, branched, feather-like structures are known
from heterodontosaurids, primitive neornithischians,[219] and
theropods,[220] 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. [11][221] Feather-like structures known
as pycnofibres have also been found in pterosaurs,[222] suggesting the
possibility that feather-like filaments may have been common in the bird
lineage and evolved before the appearance of dinosaurs
themselves.[218] 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.[223]
Archaeopteryx was the first fossil found that revealed a potential
connection between dinosaurs and birds. It is considered a transitional
fossil, in that it displays features of both groups. Brought to light just two
years after Charles Darwin's seminal On the Origin of Species (1859), its
discovery spurred the nascent debate between proponents of evolutionary
biology and creationism. This early bird is so dinosaur-like that, without a
clear impression of feathers in the surrounding rock, at least one specimen
was mistaken for the small theropod Compsognathus.[224] Since the 1990s,
a number of additional feathered dinosaurs have been found, providing
even stronger evidence of the close relationship between dinosaurs and
modern birds. Most of these specimens were unearthed in
the lagerstätte of the Yixian Formation, Liaoning, northeastern China,
which was part of an island continent during the Cretaceous. Though
feathers have been found in only a few locations, it is possible that non-
avian dinosaurs elsewhere in the world were also feathered. The lack of
widespread fossil evidence for feathered non-avian dinosaurs may be
because delicate features like skin and feathers are seldom preserved by
fossilization and thus often absent from the fossil record.[225]
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,[226][227][228] and that maniraptoran dinosaurs with
vaned feathers were not actually dinosaurs, but convergent with
dinosaurs.[217][227] 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.[229]

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. [230]

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.[231][232] 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. [195][233]

Behavioral evidence
Fossils of the troodonts Mei and Sinornithoides demonstrate that some
dinosaurs slept with their heads tucked under their arms. [234] 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.[235] 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.[236]
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. [237]

Extinction of major groups

Main article: Cretaceous–Paleogene extinction event
All non-avian dinosaurs and most lineages of birds [238] became extinct in
a mass extinction event, called the Cretaceous–Paleogene (K-Pg)
extinction event, at the end of the Cretaceous period. Above
the Cretaceous–Paleogene boundary, which has been dated to 66.038 ±
0.025 million years ago,[239] fossils of non-avian dinosaurs disappear
abruptly; the absence of dinosaur fossils was historically used to assign
rocks to the ensuing Cenozoic. The nature of the event that caused this
mass extinction has been extensively studied since the 1970s, leading to
the development of two mechanisms that are thought to have played major
roles: an extraterrestrial impact event in the Yucatán Peninsula, along
with flood basalt volcanism in India. However, the specific mechanisms of
the extinction event and the extent of its effects on dinosaurs are still areas
of ongoing research.[240] Alongside dinosaurs, many other groups of
animals became extinct: pterosaurs, marine reptiles such as mosasaurs
and plesiosaurs, several groups of mammals, ammonites (nautilus-
like mollusks), rudists (reef-building bivalves), and various groups of
marine plankton.[241][242] In all, approximately 47% of genera and 76% of
species on Earth became extinct during the K-Pg extinction event.[243] The
relatively large size of most dinosaurs and the low diversity of small-bodied
dinosaur species at the end of the Cretaceous may have contributed to
their extinction;[244] the extinction of the bird lineages that did not survive
may also have been caused by a dependence on forest habitats or a lack
of adaptations to eating seeds for survival.[245][246]

Pre-extinction diversity
Just before the K-Pg extinction event, the number of non-avian dinosaur
species that existed globally has been estimated at between 628 and
1078.[247] It remains uncertain whether the diversity of dinosaurs was in
gradual decline before the K-Pg extinction event, or whether dinosaurs
were actually thriving prior to the extinction. Rock formations from
the Maastrichtian epoch, which directly preceded the extinction, have been
found to have lower diversity than the preceding Campanian epoch, which
led to the prevailing view of a long-term decline in
diversity.[241][242][248] However, these comparisons did not account either for
varying preservation potential between rock units or for different extents of
exploration and excavation.[240] In 1984, Dale Russell carried out an
analysis to account for these biases, and found no evidence of a
decline;[249] another analysis by David Fastovsky and colleagues in 2004
even showed that dinosaur diversity continually increased until the
extinction,[250] but this analysis has been rebutted.[251] Since then, different
approaches based on statistics and mathematical models have variously
supported either a sudden extinction[240][247][252] or a gradual
decline.[253][254] End-Cretaceous trends in diversity may have varied between
dinosaur lineages: it has been suggested that sauropods were not in
decline, while ornithischians and theropods were in decline.[255][256]

Impact event
Main article: Chicxulub crater

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

The bolide impact hypothesis, first brought to wide attention in 1980

by Walter Alvarez, Luis Alvarez, and colleagues, attributes the K-Pg
extinction event to a bolide (extraterrestrial projectile) impact.[257] Alvarez
and colleagues proposed that a sudden increase in iridium levels,
recorded around the world in rock deposits at the Cretaceous–Paleogene
boundary, was direct evidence of the impact. [258] Shocked quartz, indicative
of a strong shockwave emanating from an impact, was also found
worldwide.[259] The actual impact site remained elusive until
a crater measuring 180 km (110 mi) wide was discovered in the Yucatán
Peninsula of southeastern Mexico, and was publicized in a 1991 paper
by Alan Hildebrand and colleagues.[260] Now, the bulk of the evidence
suggests that a bolide 5 to 15 kilometers (3.1 to 9.3 miles) wide impacted
the Yucatán Peninsula 66 million years ago, forming this crater[261] and
creating a "kill mechanism" that triggered the extinction event. [262][263][264]
Within hours, the Chicxulub impact would have created immediate effects
such as earthquakes,[265] tsunamis,[266] and a global firestorm that likely
killed unsheltered animals and started wildfires.[267][268] However, it would
also have had longer-term consequences for the environment. Within
days, sulphate aerosols released from rocks at the impact site would have
contributed to acid rain and ocean acidification.[269][270] Soot aerosols are
thought to have spread around the world over the ensuing months and
years; they would have cooled the surface of the Earth by
reflecting thermal radiation, and greatly slowed photosynthesis by blocking
out sunlight, thus creating an impact winter.[240][271][272] (This role was
ascribed to sulphate aerosols until experiments demonstrated
otherwise.[270]) The cessation of photosynthesis would have led to the
collapse of food webs depending on leafy plants, which included all
dinosaurs save for grain-eating birds.[246]

Deccan Traps
Main article: Deccan Traps
At the time of the K-Pg extinction, the Deccan Traps flood basalts of India
were actively erupting. The eruptions can be separated into three phases
around the K-Pg boundary, two prior to the boundary and one after. The
second phase, which occurred very close to the boundary, would have
extruded 70 to 80% of the volume of these eruptions in intermittent pulses
that occurred around 100,000 years apart.[273][274] Greenhouse gases such
as carbon dioxide and sulphur dioxide would have been released by this
volcanic activity,[275][276] resulting in climate change through temperature
perturbations of roughly 3 °C (5.4 °F) but possibly as high as 7 °C
(13 °F).[277] Like the Chicxulub impact, the eruptions may also have
released sulphate aerosols, which would have caused acid rain and global
cooling.[278] However, due to large error margins in the dating of the
eruptions, the role of the Deccan Traps in the K-Pg extinction remains
unclear.[239][240][279]
Before 2000, arguments that the Deccan Traps eruptions—as opposed to
the Chicxulub impact—caused the extinction were usually linked to the
view that the extinction was gradual. Prior to the discovery of the
Chicxulub crater, the Deccan Traps were used to explain the global iridium
layer;[275][280] even after the crater's discovery, the impact was still thought to
only have had a regional, not global, effect on the extinction event. [281] In
response, Luis Alvarez rejected volcanic activity as an explanation for the
iridium layer and the extinction as a whole. [282] Since then, however, most
researchers have adopted a more moderate position, which identifies the
Chicxulub impact as the primary progenitor of the extinction while also
recognizing that the Deccan Traps may also have played a role. Walter
Alvarez himself has acknowledged that the Deccan Traps and other
ecological factors may have contributed to the extinctions in addition to the
Chicxulub impact.[283] Some estimates have placed the start of the second
phase in the Deccan Traps eruptions within 50,000 years after the
Chicxulub impact.[284] Combined with mathematical modelling of the seismic
waves that would have been generated by the impact, this has led to the
suggestion that the Chicxulub impact may have triggered these eruptions
by increasing the permeability of the mantle plume underlying the Deccan
Traps.[285][286]
Whether the Deccan Traps were a major cause of the extinction, on par
with the Chicxulub impact, remains uncertain. Proponents consider the
climatic impact of the sulphur dioxide released to have been on par with
the Chicxulub impact, and also note the role of flood basalt volcanism in
other mass extinctions like the Permian-Triassic extinction
event.[287][288] They consider the Chicxulub impact to have worsened the
ongoing climate change caused by the eruptions. [289] Meanwhile, detractors
point out the sudden nature of the extinction and that other pulses in
Deccan Traps activity of comparable magnitude did not appear to have
caused extinctions. They also contend that the causes of different mass
extinctions should be assessed separately. [290] In 2020, Alfio Chiarenza and
colleagues suggested that the Deccan Traps may even have had the
opposite effect: they suggested that the long-term warming caused by its
carbon dioxide emissions may have dampened the impact winter from the
Chicxulub impact.[264]

Possible Paleocene survivors

Non-avian dinosaur remains have occasionally been found above the K-Pg
boundary. In 2000, Spencer Lucas and colleagues reported the discovery
of a single hadrosaur right femur in the San Juan Basin of New Mexico,
and described it as evidence of Paleocene dinosaurs. The rock unit in
which the bone was discovered has been dated to the
early Paleocene epoch, approximately 64.8 million years ago.[291] If the
bone was not re-deposited by weathering action, it would provide evidence
that some dinosaur populations may have survived at least half a million
years into the Cenozoic.[292] Other evidence includes the presence of
dinosaur remains in the Hell Creek Formation up to 1.3 m (4.3 ft) above
the Cretaceous–Paleogene boundary, representing 40,000 years of
elapsed time. This has been used to support the view that the K-Pg
extinction was gradual.[293] However, these supposed Paleocene dinosaurs
are considered by many other researchers to be reworked, that is, washed
out of their original locations and then re-buried in younger
sediments.[294][295][296] The age estimates have also been considered
unreliable.[297]

Cultural depictions
Main article: Cultural depictions of dinosaurs

Outdated Iguanodon statues created by Benjamin Waterhouse Hawkins for

the Crystal Palace Park in 1853

The battles that may have occurred

between Tyrannosaurus and Triceratops are a recurring theme in popular
science and dinosaurs' depiction in culture

By human standards, dinosaurs were creatures of fantastic appearance

and often enormous size. As such, they have captured the popular
imagination and become an enduring part of human culture. The entry of
the word "dinosaur" into the common vernacular reflects the animals'
cultural importance: in English, "dinosaur" is commonly used to describe
anything that is impractically large, obsolete, or bound for extinction. [298]
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 was 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.[299] The enduring popularity of dinosaurs, in its turn, has resulted in
significant public funding for dinosaur science, and has frequently spurred
new discoveries. In the United States, for example, the competition
between museums for public attention led directly to the Bone Wars of the
1880s and 1890s, during which a pair of feuding paleontologists made
enormous scientific contributions.[300]
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,[301] 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 1914 animated film Gertie the
Dinosaur (featuring the first animated dinosaur), the iconic 1933 film King
Kong, the 1954 Godzilla and its many sequels, the best-selling 1990
novel Jurassic Park by Michael Crichton and its 1993 film adaptation are
just a few notable examples of dinosaur appearances in fiction. Authors of
general-interest non-fiction works about dinosaurs, including some
prominent paleontologists, have often sought to use the animals as a way
to educate readers about science in general. Dinosaurs are ubiquitous
in advertising; numerous companies have referenced dinosaurs in printed
or televised advertisements, either in order to sell their own products or in
order to characterize their rivals as slow-moving, dim-witted, or
obsolete.[302][303]