Dinosaurs In Amusement Parks
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Dinosaurs are a diverse group of
reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
s of the
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
Dinosauria. They first appeared during the
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
period Period may refer to: Common uses * Era, a length or span of time * Full stop (or period), a punctuation mark Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (or rhetorical period), a concept ...
, between 243 and 233.23 
million years ago The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago). ...
(mya), although the exact origin and timing of the
evolution of dinosaurs Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
is the subject of active research. They became the dominant terrestrial
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
s after the
Triassic–Jurassic extinction event The Triassic–Jurassic (Tr-J) extinction event, often called the end-Triassic extinction, marks the boundary between the Triassic and Jurassic periods, , and is one of the top five major extinction events of the Phanerozoic eon, profoundly affect ...
201.3 mya; their dominance continued throughout the
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
and
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
periods. The
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
record shows that
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...
s are
feathered dinosaur A feathered dinosaur is any species of dinosaur possessing feathers. While this includes all species of birds, there is a hypothesis that many, if not all non-avian dinosaur species also possessed feathers in some shape or form. It has been su ...
s, having evolved from earlier
theropods Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
during the
Late Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
epoch, and are the only dinosaur lineage known to have survived the
Cretaceous–Paleogene extinction event The Cretaceous–Paleogene (K–Pg) extinction event (also known as the Cretaceous–Tertiary extinction) was a sudden mass extinction of three-quarters of the plant and animal species on Earth, approximately 66 million years ago. With the ...
approximately 66 mya. Dinosaurs can therefore be divided into avian dinosaurs—birds—and the extinct non-avian dinosaurs, which are all dinosaurs other than birds. Dinosaurs are varied from
taxonomic Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
, morphological and
ecological Ecology () is the study of the relationships between living organisms, including humans, and their physical environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere level. Ecology overlaps wi ...
standpoints. Birds, at over 10,700 living
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
, are among the most diverse group of vertebrates. Using fossil evidence,
paleontologists Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
have identified over 900 distinct
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
and more than 1,000 different species of non-avian dinosaurs. Dinosaurs are represented on every continent by both
extant Extant is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Extant taxon, a taxon which is not extinct, ...
species (birds) and fossil remains. Through the first half of the 20th century, before birds were recognized as dinosaurs, most of the scientific community believed dinosaurs to have been sluggish and cold-blooded. Most research conducted since the 1970s, however, has indicated that dinosaurs were active animals with elevated
metabolism Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run cell ...
s and numerous adaptations for social interaction. Some were
herbivorous A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
, others
carnivorous A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other sof ...
. Evidence suggests that all dinosaurs were egg-laying, and that
nest A nest is a structure built for certain animals to hold eggs or young. Although nests are most closely associated with birds, members of all classes of vertebrates and some invertebrates construct nests. They may be composed of organic materia ...
-building was a trait shared by many dinosaurs, both avian and non-avian. While dinosaurs were ancestrally
bipedal Bipedalism is a form of terrestrial locomotion where an organism moves by means of its two rear limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' 'double' ...
, many
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
groups included
quadrupedal Quadrupedalism is a form of locomotion where four limbs are used to bear weight and move around. An animal or machine that usually maintains a four-legged posture and moves using all four limbs is said to be a quadruped (from Latin ''quattuor' ...
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 A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
modifications such as bony armor and spines. While the dinosaurs' modern-day surviving avian lineage (birds) are generally small due to the constraints of flight, many prehistoric dinosaurs (non-avian and avian) were large-bodied—the largest
sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their bo ...
dinosaurs are estimated to have reached lengths of and heights of and were the largest land animals of all time. The misconception that non-avian dinosaurs were uniformly gigantic is based in part on
preservation bias Taphonomy is the study of how organisms decay and become fossilized or preserved in the paleontological record. The term ''taphonomy'' (from Greek , 'burial' and , 'law') was introduced to paleontology in 1940 by Soviet scientist Ivan Efremov t ...
, as large, sturdy
bone A bone is a Stiffness, rigid Organ (biology), organ that constitutes part of the skeleton in most vertebrate animals. Bones protect the various other organs of the body, produce red blood cell, red and white blood cells, store minerals, provid ...
s are more likely to last until they are fossilized. Many dinosaurs were quite small, some measuring about in length. The first dinosaur fossils were recognized in the early 19th century, with the name "dinosaur" (meaning "terrible lizard") being coined by Sir
Richard Owen Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomist and paleontologist. Owen is generally considered to have been an outstanding naturalist with a remarkable gift for interpreting fossils. Owe ...
in 1841 to refer to these "great fossil lizards". Since then, mounted fossil dinosaur skeletons have been major attractions at museums worldwide, and dinosaurs have become an enduring part of popular culture. The large sizes of some dinosaurs, as well as their seemingly monstrous and fantastic nature, have ensured their regular appearance in best-selling books and films, such as ''
Jurassic Park ''Jurassic Park'', later also referred to as ''Jurassic World'', is an American science fiction media franchise created by Michael Crichton and centered on a disastrous attempt to create a theme park of cloned dinosaurs. It began in 1990 when ...
''. Persistent public enthusiasm for the animals has resulted in significant funding for dinosaur science, and new discoveries are regularly covered by the media.


Definition

Under
phylogenetic nomenclature Phylogenetic nomenclature is a method of nomenclature for taxa in biology that uses phylogenetic definitions for taxon names as explained below. This contrasts with the traditional approach, in which taxon names are defined by a '' type'', which ...
, dinosaurs are usually defined as the group consisting of the
most recent common ancestor In biology and genetic genealogy, the most recent common ancestor (MRCA), also known as the last common ancestor (LCA) or concestor, of a set of organisms is the most recent individual from which all the organisms of the set are descended. The ...
(MRCA) of ''
Triceratops ''Triceratops'' ( ; ) is a genus of herbivore, herbivorous Chasmosaurinae, chasmosaurine Ceratopsidae, ceratopsid dinosaur that first appeared during the late Maastrichtian stage of the Late Cretaceous Period (geology), period, about 68 m ...
'' and
modern birds Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
(Neornithes), and all its descendants. It has also been suggested that Dinosauria be defined with respect to the MRCA of ''
Megalosaurus ''Megalosaurus'' (meaning "great lizard", from Greek , ', meaning 'big', 'tall' or 'great' and , ', meaning 'lizard') is an extinct genus of large carnivorous theropod dinosaurs of the Middle Jurassic period (Bathonian stage, 166 million years a ...
'' and ''
Iguanodon ''Iguanodon'' ( ; meaning 'iguana-tooth'), named in 1825, is a genus of iguanodontian dinosaur. While many species have been classified in the genus ''Iguanodon'', dating from the late Jurassic Period to the early Cretaceous Period of Asia, Eu ...
'', because these were two of the three genera cited by
Richard Owen Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomist and paleontologist. Owen is generally considered to have been an outstanding naturalist with a remarkable gift for interpreting fossils. Owe ...
when he recognized the Dinosauria. Both definitions result in the same set of animals being defined as dinosaurs: "Dinosauria =
Ornithischia Ornithischia () is an extinct order of mainly herbivorous dinosaurs characterized by a pelvic structure superficially similar to that of birds. The name ''Ornithischia'', or "bird-hipped", reflects this similarity and is derived from the Greek s ...
+
Saurischia Saurischia ( , meaning "reptile-hipped" from the Greek ' () meaning 'lizard' and ' () meaning 'hip joint') is one of the two basic divisions of dinosaurs (the other being Ornithischia), classified by their hip structure. Saurischia and Ornithis ...
". This definition includes major groups such as
ankylosauria Ankylosauria is a group of herbivorous dinosaurs of the order Ornithischia. It includes the great majority of dinosaurs with armor in the form of bony osteoderms, similar to turtles. Ankylosaurs were bulky quadrupeds, with short, powerful limbs. ...
ns (armored herbivorous quadrupeds),
stegosauria Stegosauria is a group of herbivorous ornithischian dinosaurs that lived during the Jurassic and early Cretaceous periods. Stegosaurian fossils have been found mostly in the Northern Hemisphere, predominantly in what is now North America, Europe, ...
ns (plated herbivorous quadrupeds),
ceratopsia Ceratopsia or Ceratopia ( or ; Greek: "horned faces") is a group of herbivorous, beaked dinosaurs that thrived in what are now North America, Europe, and Asia, during the Cretaceous Period, although ancestral forms lived earlier, in the Jurassic. ...
ns (bipedal or quadrupedal herbivores with
neck frill A neck frill is the relatively extensive margin seen on the back of the heads of reptiles with either a bony support such as those present on the skulls of dinosaurs of the suborder Marginocephalia or a cartilaginous one as in the frill-necked ...
s),
pachycephalosauria Pachycephalosauria (; from Greek παχυκεφαλόσαυρος for 'thick headed lizards') is a clade of ornithischian dinosaurs. Along with Ceratopsia, it makes up the clade Marginocephalia. With the exception of two species, most pachycephal ...
ns (bipedal herbivores with thick skulls),
ornithopod Ornithopoda () is a clade of ornithischian dinosaurs, called ornithopods (), that started out as small, bipedal running grazers and grew in size and numbers until they became one of the most successful groups of herbivores in the Cretaceous world ...
s (bipedal or quadrupedal herbivores including " duck-bills"),
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
s (mostly bipedal carnivores and birds), and
sauropodomorphs Sauropodomorpha ( ; from Greek, meaning "lizard-footed forms") is an extinct clade of long-necked, herbivorous, saurischian dinosaurs that includes the sauropods and their ancestral relatives. Sauropods generally grew to very large sizes, had lon ...
(mostly large herbivorous quadrupeds with long necks and tails). Birds are now recognized as being the sole surviving lineage of theropod dinosaurs. In traditional
taxonomy Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
, birds were considered a separate
class Class or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or objects * Class (philosophy), an analytical concept used differentl ...
that had evolved from dinosaurs, a distinct
superorder Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...
. However, a majority of contemporary paleontologists concerned with dinosaurs reject the traditional style of classification in favor of
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
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 Maniraptora is a clade of coelurosaurian dinosaurs which includes the birds and the non-avian dinosaurs that were more closely related to them than to ''Ornithomimus velox''. It contains the major subgroups Avialae, Deinonychosauria, Oviraptoros ...
, which are
coelurosaurs Coelurosauria (; from Greek, meaning "hollow tailed lizards") is the clade containing all theropod dinosaurs more closely related to birds than to carnosaurs. Coelurosauria is a subgroup of theropod dinosaurs that includes compsognathids, tyrann ...
, which are theropods, which are saurischians, which are dinosaurs. 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 Ornithoscelida () is a proposed clade that includes various major groupings of dinosaurs. An order Ornithoscelida was originally proposed by Thomas Henry Huxley but later abandoned in favor of Harry Govier Seeley's division of Dinosauria into Sau ...
to refer to the group containing Ornithischia and Theropoda. Dinosauria itself was re-defined as the last common ancestor of ''Triceratops horridus'', ''
Passer domesticus The house sparrow (''Passer domesticus'') is a bird of the sparrow family Passeridae, found in most parts of the world. It is a small bird that has a typical length of and a mass of . Females and young birds are coloured pale brown and grey, an ...
'' and ''
Diplodocus carnegii ''Diplodocus'' (, , or ) was a genus of diplodocid sauropod dinosaurs, whose fossils were first discovered in 1877 by S. W. Williston. The generic name, coined by Othniel Charles Marsh in 1878, is a neo-Latin term derived from Greek δι ...
'', and all of its descendants, to ensure that sauropods and kin remain included as dinosaurs. "A version of this article appears in print on March 28, 2017, on Page D6 of the New York edition with the headline: Shaking Up the Dinosaur Family Tree." "This file contains Supplementary Text and Data, Supplementary Tables 1-3 and additional references."
Supplementary Information
/ref>


General description

Using one of the above definitions, dinosaurs can be generally described as
archosaur Archosauria () is a clade of diapsids, with birds and crocodilians as the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term which includes birds. Extinct archosaurs include non-avian d ...
s with hind limbs held erect beneath the body. Other prehistoric animals, including
pterosaur Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 to ...
s,
mosasaur Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Greek ' meaning 'lizard') comprise a group of extinct, large marine reptiles from the Late Cretaceous. Their first fossil remains were discovered in a limestone quarry at Maastricht on th ...
s,
ichthyosaur Ichthyosaurs (Ancient Greek for "fish lizard" – and ) are large extinct marine reptiles. Ichthyosaurs belong to the order known as Ichthyosauria or Ichthyopterygia ('fish flippers' – a designation introduced by Sir Richard Owen in 1842, altho ...
s,
plesiosaurs The Plesiosauria (; Greek: πλησίος, ''plesios'', meaning "near to" and ''sauros'', meaning "lizard") or plesiosaurs are an order or clade of extinct Mesozoic marine reptiles, belonging to the Sauropterygia. Plesiosaurs first appeared ...
, and ''
Dimetrodon ''Dimetrodon'' ( or ,) meaning "two measures of teeth,” is an extinct genus of non-mammalian synapsid that lived during the Cisuralian (Early Permian), around 295–272 million years ago (Mya). It is a member of the family Sphenacodontid ...
'', while often popularly conceived of as dinosaurs, are not taxonomically classified as dinosaurs. Pterosaurs are distantly related to dinosaurs, being members of the clade
Ornithodira Avemetatarsalia (meaning "bird metatarsals") is a clade of diapsid reptiles containing all archosaurs more closely related to birds than to crocodilians. The two most successful groups of avemetatarsalians were the dinosaurs and pterosaurs. Dinos ...
. The other groups mentioned are, like dinosaurs and pterosaurs, members of
Sauropsida Sauropsida ("lizard faces") is a clade of amniotes, broadly equivalent to the class Reptilia. Sauropsida is the sister taxon to Synapsida, the other clade of amniotes which includes mammals as its only modern representatives. Although early syna ...
(the reptile and bird clade), except ''Dimetrodon'' (which is a
synapsid Synapsids + (, 'arch') > () "having a fused arch"; synonymous with ''theropsids'' (Greek, "beast-face") are one of the two major groups of animals that evolved from basal amniotes, the other being the sauropsids, the group that includes reptil ...
). None of them had the erect hind limb posture characteristic of true dinosaurs. Dinosaurs were the dominant terrestrial vertebrates of the
Mesozoic The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceo ...
Era An era is a span of time defined for the purposes of chronology or historiography, as in the regnal eras in the history of a given monarchy, a calendar era used for a given calendar, or the geological eras defined for the history of Earth. Comp ...
, especially the Jurassic and Cretaceous periods. Other groups of animals were restricted in size and niches;
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
s, for example, rarely exceeded the size of a domestic cat, and were generally rodent-sized carnivores of small prey. 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. A 1995 study predicted that about 3,400 dinosaur genera ever existed, including many that would not have been preserved in the fossil record. In 2016, the estimated number of dinosaur species that existed in the Mesozoic was 1,543–2,468. In 2021, the number of modern-day birds (avian dinosaurs) was estimated to be at 10,806 species. 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 ''Anchisaurus'' is a genus of basal sauropodomorph dinosaur. It lived during the Early Jurassic Period, and its fossils have been found in the red sandstone of the Portland Formation, Northeastern United States, which was deposited from the He ...
'' 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 The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic-J ...
epoch. Modern birds inhabit most available habitats, from terrestrial to marine, and there is evidence that some non-avian dinosaurs (such as ''
Microraptor ''Microraptor'' (Greek, μικρός, ''mīkros'': "small"; Latin, ''raptor'': "one who seizes") is a genus of small, four-winged dromaeosaurid dinosaurs. Numerous well-preserved fossil specimens have been recovered from Liaoning, China. They dat ...
'') could fly or at least glide, and others, such as
spinosaurids The Spinosauridae (or spinosaurids) are a clade or Family (taxonomy), family of tetanuran theropod dinosaurs comprising ten to seventeen known genera. They came into prominence during the Cretaceous Geological period, period. Spinosaurid fossils h ...
, had
semiaquatic In biology, semiaquatic can refer to various types of animals that spend part of their time in water, or plants that naturally grow partially submerged in water. Examples are given below. Semiaquatic animals Semiaquatic animals include: * Verte ...
habits.


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 In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
of such a group. A detailed assessment of archosaur interrelations by
Sterling Nesbitt Sterling Nesbitt (born March 25, 1982, in Mesa, Arizona) is an American paleontologist best known for his work on the origin and early evolutionary patterns of archosaurs. He is currently an associate professor at Virginia Tech in the Department of ...
confirmed or found the following twelve unambiguous synapomorphies, some previously known: * In the
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...
, a supratemporal fossa (excavation) is present in front of the
supratemporal fenestra The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...
, the main opening in the rear skull roof *
Epipophyses Epipophyses are bony projections of the cervical vertebrae found in archosauromorphs, particularly dinosaurs (including some basal birds). These paired processes sit above the postzygapophyses on the rear of the vertebral neural arch. Their morp ...
, obliquely backward-pointing processes on the rear top corners of the anterior (front) neck
vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...
e behind the
atlas An atlas is a collection of maps; it is typically a bundle of maps of Earth or of a region of Earth. Atlases have traditionally been bound into book form, but today many atlases are in multimedia formats. In addition to presenting geographic ...
and
axis An axis (plural ''axes'') is an imaginary line around which an object rotates or is symmetrical. Axis may also refer to: Mathematics * Axis of rotation: see rotation around a fixed axis *Axis (mathematics), a designator for a Cartesian-coordinate ...
, 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 The humerus (; ) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of a roun ...
(upper arm bone) *
Radius In classical geometry, a radius ( : radii) of a circle or sphere is any of the line segments from its center to its perimeter, and in more modern usage, it is also their length. The name comes from the latin ''radius'', meaning ray but also the ...
, a lower arm bone, shorter than 80% of humerus length *
Fourth trochanter The fourth trochanter is a shared characteristic common to archosaurs. It is a knob-like feature on the posterior-medial side of the middle of the femur shaft that serves as a muscle attachment, mainly for the ''musculus caudofemoralis longus'', ...
(projection where the
caudofemoralis The caudofemoralis (from the Latin ''cauda'', tail and ''femur'', thighbone) is a muscle found in the pelvic limb of mostly all animals possessing a tail. It is thus found in nearly all tetrapods. Location The caudofemoralis spans plesiomorphi ...
muscle attaches on the inner rear shaft) on the
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates with ...
(thigh bone) is a sharp flange * Fourth trochanter asymmetrical, with distal, lower, margin forming a steeper angle to the shaft * On the
astragalus ''Astragalus'' is a large genus of over 3,000 species of herbs and small shrubs, belonging to the legume family Fabaceae and the subfamily Faboideae. It is the largest genus of plants in terms of described species. The genus is native to tempe ...
and
calcaneum In humans and many other primates, the calcaneus (; from the Latin ''calcaneus'' or ''calcaneum'', meaning heel) or heel bone is a bone of the tarsus of the foot which constitutes the heel. In some other animals, it is the point of the hock. St ...
, upper ankle bones, the proximal articular facet, the top connecting surface, for the
fibula The fibula or calf bone is a leg bone on the lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long bones. Its upper extremity is ...
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 The ischium () form ...
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 The cnemial crest is a crestlike prominence located at the front side of the head of the tibiotarsus or tibia in the legs of many mammals and reptiles (including birds and other dinosaurs). The main extensor muscle of the thigh In human anatomy ...
on the
tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...
(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. 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
diapsid Diapsids ("two arches") are a clade of sauropsids, distinguished from more primitive eureptiles by the presence of two holes, known as temporal fenestrae, in each side of their skulls. The group first appeared about three hundred million years ago ...
s, dinosaurs ancestrally had two pairs of
Infratemporal fenestra An infratemporal fenestra, also called the lateral temporal fenestra or simply temporal fenestra, is an opening in the skull behind the orbit in some animals. It is ventrally bordered by a zygomatic arch. An opening in front of the eye sockets ...
e (openings in the skull behind the eyes), and as members of the diapsid group Archosauria, had additional openings in the
snout A snout is the protruding portion of an animal's face, consisting of its nose, mouth, and jaw. In many animals, the structure is called a muzzle, rostrum, or proboscis. The wet furless surface around the nostrils of the nose of many mammals is c ...
and lower jaw. 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 The scapula (plural scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on eithe ...
, or shoulder blade; a
sacrum The sacrum (plural: ''sacra'' or ''sacrums''), in human anatomy, is a large, triangular bone at the base of the spine that forms by the fusing of the sacral vertebrae (S1S5) between ages 18 and 30. The sacrum situates at the upper, back part ...
composed of three or more fused vertebrae (three are found in some other archosaurs, but only two are found in ''
Herrerasaurus ''Herrerasaurus'' is a genus of saurischian dinosaur from the Late Triassic period. This genus was one of the earliest dinosaurs from the fossil record. Its name means "Herrera's lizard", after the rancher who discovered the first specimen in ...
''); and a perforate
acetabulum The acetabulum (), also called the cotyloid cavity, is a concave surface of the pelvis. The head of the femur meets with the pelvis at the acetabulum, forming the hip joint. Structure There are three bones of the ''os coxae'' (hip bone) that c ...
, or hip socket, with a hole at the center of its inside surface (closed in ''
Saturnalia tupiniquim ''Saturnalia'' is an extinct genus of basal sauropodomorph dinosaur known from the Late Triassic Santa Maria Formation of Rio Grande do Sul, southern Brazil and possibly the Pebbly Arkose Formation, Zimbabwe. Discovery and naming ''Saturnal ...
'', for example). Another difficulty of determining distinctly dinosaurian features is that early dinosaurs and other archosaurs from the
Late Triassic The Late Triassic is the third and final epoch (geology), epoch of the Triassic geologic time scale, Period in the geologic time scale, spanning the time between annum, Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch ...
epoch are often poorly known and were similar in many ways; these animals have sometimes been misidentified in the literature. 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. 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. Their erect posture enabled early dinosaurs to breathe easily while moving, which likely permitted stamina and activity levels that surpassed those of "sprawling" reptiles. Erect limbs probably also helped support the
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
of large size by reducing bending stresses on limbs. Some non-dinosaurian archosaurs, including
rauisuchia "Rauisuchia" is a paraphyletic group of mostly large and carnivorous Triassic archosaurs. Rauisuchians are a category of archosaurs within a larger group called Pseudosuchia, which encompasses all archosaurs more closely related to crocodilia ...
ns, 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.


History of study


Pre-scientific history

Dinosaur fossils have been known for millennia, although their true nature was not recognized. The Chinese considered them to be
dragon A dragon is a reptilian legendary creature that appears in the folklore of many cultures worldwide. Beliefs about dragons vary considerably through regions, but dragons in western cultures since the High Middle Ages have often been depicted as ...
bones and documented them as such. For example, '' Huayang Guo Zhi'' (), a
gazetteer A gazetteer is a geographical index or directory used in conjunction with a map or atlas.Aurousseau, 61. It typically contains information concerning the geographical makeup, social statistics and physical features of a country, region, or co ...
compiled by
Chang Qu Chang Qu () (c. 291 – c. 361 CE) was a 4th-century Chinese historian of the Cheng Han dynasty, who wrote the ''Chronicles of Huayang'' or ''Records of the States South of Mount Hua Mount Hua () is a mountain located near the city of Huayin ...
 () during the
Western Jin Dynasty Western may refer to: Places *Western, Nebraska, a village in the US *Western, New York, a town in the US *Western Creek, Tasmania, a locality in Australia *Western Junction, Tasmania, a locality in Australia *Western world, countries that id ...
(265–316), reported the discovery of dragon bones at Wucheng in
Sichuan Sichuan (; zh, c=, labels=no, ; zh, p=Sìchuān; alternatively romanized as Szechuan or Szechwan; formerly also referred to as "West China" or "Western China" by Protestant missions) is a province in Southwest China occupying most of the ...
Province. Villagers in central
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
have long unearthed fossilized "dragon bones" for use in
traditional medicines Traditional medicine (also known as indigenous medicine or folk medicine) comprises medical aspects of traditional knowledge that developed over generations within the folk beliefs of various societies, including indigenous peoples, before the ...
. In
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
, dinosaur fossils were generally believed to be the remains of
giant In folklore, giants (from Ancient Greek: '' gigas'', cognate giga-) are beings of human-like appearance, but are at times prodigious in size and strength or bear an otherwise notable appearance. The word ''giant'' is first attested in 1297 fr ...
s and other
biblical The Bible (from Koine Greek , , 'the books') is a collection of religious texts or scriptures that are held to be sacred in Christianity, Judaism, Samaritanism, and many other religions. The Bible is an anthologya compilation of texts of a ...
creatures.


Early dinosaur research

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'', was recovered from a limestone quarry at
Cornwell Cornwell can refer to: People * Almon Cornwell (1820–1893), American farmer-politician in Wisconsin * Anita Cornwell (1923–2023), American author * Bernard Cornwell (born 1944), British historical novelist * Charlotte Cornwell (1949–2021), Bri ...
near
Chipping Norton Chipping Norton is a market town and civil parish in the Cotswold Hills in the West Oxfordshire district of Oxfordshire, England, about south-west of Banbury and north-west of Oxford. The 2011 Census recorded the civil parish population as ...
, Oxfordshire, in 1676. The fragment was sent to
Robert Plot Robert Plot (13 December 1640 – 30 April 1696) was an English naturalist, first Professor of Chemistry at the University of Oxford, and the first keeper of the Ashmolean Museum. Early life and education Born in Borden, Kent to parents Robe ...
, Professor of Chemistry at the
University of Oxford , mottoeng = The Lord is my light , established = , endowment = £6.1 billion (including colleges) (2019) , budget = £2.145 billion (2019–20) , chancellor ...
and first curator of the
Ashmolean Museum The Ashmolean Museum of Art and Archaeology () on Beaumont Street, Oxford, England, is Britain's first public museum. Its first building was erected in 1678–1683 to house the cabinet of curiosities that Elias Ashmole gave to the University of ...
, who published a description in his ''The Natural History of Oxford-shire'' (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 femur of a huge human, perhaps a Titan or another type of giant featured in legends.
Edward Lhuyd Edward Lhuyd FRS (; occasionally written Llwyd in line with modern Welsh orthography, 1660 – 30 June 1709) was a Welsh naturalist, botanist, linguist, geographer and antiquary. He is also named in a Latinate form as Eduardus Luidius. Life ...
, a friend of
Sir Isaac Newton Sir Isaac Newton (25 December 1642 – 20 March 1726/27) was an English mathematician, physicist, astronomer, alchemist, theologian, and author (described in his time as a "natural philosopher"), widely recognised as one of the great ...
, 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, Rutellum impicatum", that had been found in Caswell, near Witney, Oxfordshire. 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. 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. 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. The term is derived . 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. 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 Albert, Prince Consort, 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.


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, Hadrosaurus foulkii''. It was an extremely important find: ''Hadrosaurus'' was one of the first nearly complete dinosaur skeletons found (Iguanodon#Gideon Mantell, Sir Richard Owen, and the discovery of dinosaurs, the first was in 1834, in Maidstone, 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. 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.


"Dinosaur renaissance" and beyond

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. 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.


Soft tissue and DNA

One of the best examples of soft-tissue impressions in a fossil dinosaur was discovered in the Pietraroja Formation, Pietraroja Plattenkalk in southern Italy. The discovery was reported in 1998, and described the specimen of a small, juvenile coelurosaur, ''Scipionyx, Scipionyx samniticus''. The fossil includes portions of the intestines, colon, liver, muscles, and windpipe of this dinosaur. In the March 2005 issue of ''Science (journal), 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. 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, Osteon, 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. 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, 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. The extraction of ancient DNA from dinosaur fossils has been reported on two separate occasions; upon further inspection and peer review, however, neither of these reports could be confirmed. 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. In addition, several proteins, including hemoglobin, have putatively been detected in dinosaur fossils. 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.


Evolutionary history


Origins and early evolution

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. Radiometric dating of the Ischigualasto Formation of Argentina where the early dinosaur genus ''Eoraptor'' was found date it as 231.4 million years old. ''Eoraptor'' is thought to resemble the Common descent, common ancestor of all dinosaurs; if this is true, its traits suggest that the first dinosaurs were small, bipedal predation, predators. 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, 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. 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''. When dinosaurs appeared, they were not the dominant terrestrial animals. The terrestrial habitats were occupied by various types of Archosauromorpha, archosauromorphs and therapsids, like cynodonts and rhynchosaurs. Their main competitors were the pseudosuchians, such as aetosaurs, ornithosuchidae, ornithosuchids and rauisuchians, which were more successful than the dinosaurs. 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 (phylogenetics), basal archosauromorphs, including the Protorosauria, 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 Stage (stratigraphy), stages, respectively, and the exact date of their extinction is uncertain. These losses left behind a land fauna of crocodylomorpha, crocodylomorphs, dinosaurs, mammals, pterosaurians, and turtles. The first few lines of early dinosaurs Adaptive radiation, diversified through the Carnian and Norian stages of the Triassic, possibly by occupying the niches of the groups that became extinct. Also notably, there was a heightened rate of extinction during the Carnian pluvial episode, Carnian pluvial event.


Evolution and paleobiogeography

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 coelophysoidea, coelophysoid carnivores and early sauropodomorph herbivores. Gymnosperm plants (particularly Pinophyta, conifers), a potential food source, Evolutionary radiation, 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. The general homogeneity of dinosaurian faunas continued into the Middle and Late Jurassic, where most localities had predators consisting of ceratosaurians, Megalosauroidea, megalosauroids, and Allosauroidea, allosauroids, and herbivores consisting of stegosaurian ornithischians and large sauropods. Examples of this include the Morrison Formation of North America and Tendaguru Formation, Tendaguru Beds of Tanzania. Dinosaurs in China show some differences, with specialized Metriacanthosauridae, metriacanthosaurid theropods and unusual, long-necked sauropods like ''Mamenchisaurus''. 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. Another notable evolutionary event of the Jurassic was the appearance of true birds, descended from maniraptoran coelurosaurians. 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 brachiosauridae, brachiosaurids through Europe, North America, and northern Africa. These were later supplemented or replaced in Africa by large spinosaurid and carcharodontosauridae, carcharodontosaurid theropods, and rebbachisauridae, rebbachisaurid and titanosaurian sauropods, also found in South America. In Asia, maniraptoran coelurosaurians like dromaeosauridae, dromaeosaurids, troodontidae, troodontids, and oviraptorosaurians became the common theropods, and ankylosauridae, ankylosaurids and early ceratopsians like ''Psittacosaurus'' became important herbivores. Meanwhile, Australia was home to a fauna of basal ankylosaurians, hypsilophodonts, and iguanodontians. 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 Dinosaur tooth#Dental batteries, dental batteries, taken to its extreme in hadrosaurids. Some sauropods also evolved tooth batteries, best exemplified by the rebbachisaurid ''Nigersaurus''. There were three general dinosaur faunas in the Late Cretaceous. In the northern continents of North America and Asia, the major theropods were tyrannosauridae, 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, abelisauridae, abelisaurids were the common theropods, and titanosaurian sauropods the common herbivores. Finally, in Europe, dromaeosaurids, rhabdodontidae, rhabdodontid iguanodontians, nodosauridae, nodosaurid ankylosaurians, and titanosaurian sauropods were prevalent. Flowering plants were greatly radiating, with the first grasses appearing by the end of the Cretaceous. Grinding hadrosaurids and shearing ceratopsians became very diverse across North America and Asia. Theropods were also radiating as herbivores or omnivores, with Therizinosauria, therizinosaurians and ornithomimosaurians becoming common. 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, Dyrosauridae, dyrosaurs, sebecosuchians, turtles, lizards, snakes, Rhynchocephalia, sphenodontians, and choristoderans, also survived the event. The surviving lineages of neornithine birds, including the ancestors of modern ratites, Fowl, ducks and chickens, and a variety of Charadriiformes, 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'', Eogruidae, eogruiids, bathornithids, ratites, Geranoididae, 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. Terror birds and bathornithids occupied carnivorous guilds alongside predatory mammals, 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.


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. 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 taxon, taxa sharing a more recent common ancestor with ''Triceratops'' than with Saurischia. Anatomically, these two groups can be distinguished most noticeably by their pelvis, pelvic structure. Early saurischians—"lizard-hipped", from the Ancient Greek, Greek ' () meaning "lizard" and ' () meaning "hip joint"—retained the hip structure of their ancestors, with a pubis bone directed anatomical terms of location, cranially, or forward. This basic form was modified by rotating the pubis backward to varying degrees in several groups (''Herrerasaurus'', therizinosauroids, dromaeosaurids, and birds). Saurischia includes the theropods (exclusively bipedal and with a wide variety of diets) and sauropodomorphs (long-necked herbivores which include advanced, quadrupedal groups). 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".


Taxonomy

The following is a simplified classification of dinosaur groups based on their evolutionary relationships, and those of the main dinosaur groups Theropoda, Sauropodomorpha and Ornithischia, compiled by Justin Tweet. Further details and other hypotheses of classification may be found on individual articles. *Dinosauria :*†
Ornithischia Ornithischia () is an extinct order of mainly herbivorous dinosaurs characterized by a pelvic structure superficially similar to that of birds. The name ''Ornithischia'', or "bird-hipped", reflects this similarity and is derived from the Greek s ...
("bird-hipped"; diverse bipedal and quadrupedal herbivores) ::*†Heterodontosauridae (small herbivores/omnivores with prominent canine tooth, canine-like teeth) :::*†Genasauria ("cheeked lizards") ::::*†Thyreophora (armored dinosaurs; bipeds and quadrupeds) :::::*†Eurypoda (heavy, quadrupedal thyreophorans) ::::::*†Stegosauria (spikes and plates as primary armor) :::::::*†Huayangosauridae (small stegosaurs with flank osteoderms and tail clubs) :::::::*†Stegosauridae (large stegosaurs) ::::::*†Ankylosauria (scutes as primary armor) :::::::*†Parankylosauria (small, southern ankylosaurs with macuahuitl-like tails) :::::::*†Nodosauridae (mostly spiky, club-less ankylosaurs) :::::::*†Ankylosauridae (characterized by flat scutes) ::::::::*†Ankylosaurinae (club-tailed ankylosaurids) ::::*†Neornithischia ("new ornithischians") :::::*†Cerapoda ("horned feet") ::::::*†Marginocephalia (characterized by a cranial growth) :::::::*†Pachycephalosauria (bipeds with domed or knobby growth on skulls) :::::::*†Ceratopsia (bipeds and quadrupeds; many had neck frills and horns) ::::::::*†Chaoyangsauridae (small, frill-less basal ceratopsians) ::::::::*†Neoceratopsia ("new ceratopsians") :::::::::*†Leptoceratopsidae (little to no frills, hornless, with robust jaws) :::::::::*†Protoceratopsidae (basal ceratopsians with small frills and stubby horns) ::::::::::*†Ceratopsoidea (large-horned ceratopsians) :::::::::::*†Ceratopsidae (large, elaborately ornamented ceratopsians) ::::::::::::*†Chasmosaurinae (ceratopsids with enlarged brow horns) :::::::::::::*†Triceratopsini (very large chasmosaurines with long brow horns) ::::::::::::*†Centrosaurinae (ceratopsids mostly characterized by frill and nasal ornamentation) :::::::::::::*†Nasutoceratopsini (centrosaurines with enlarged nasal cavities) :::::::::::::*†Centrosaurini (centrosaurines with enlarged nasal horns) :::::::::::::*†Pachyrhinosaurini (mostly had nasal bosses instead of horns) ::::::*†Ornithopoda (various sizes; bipeds and quadrupeds; evolved a method of chewing using skull flexibility and numerous teeth) :::::::*†Jeholosauridae (small Asian neornithischians) :::::::*†Thescelosauridae ("wondrous lizards") ::::::::*†Orodrominae (burrowers) ::::::::*†Thescelosaurinae (large thescelosaurids) :::::::*†Iguanodontia ("iguana teeth"; advanced ornithopods) ::::::::*†Elasmaria (mostly southern ornithopods with mineralized plates along the ribs; may be thescelosaurids) ::::::::*†Rhabdodontomorpha (with distinctive dentition) :::::::::*†Rhabdodontidae (European rhabdodontomorphs) ::::::::*†Dryosauridae (mid-sized, small headed) ::::::::*†Camptosauridae (mid-sized, stocky) :::::::::*†Styracosterna ("spiked Sternum, sterna") ::::::::::*†Hadrosauriformes (ancestrally had a thumb spike) :::::::::::*†Hadrosauroidea (large quadrupedal herbivores, with teeth merged into dental batteries) ::::::::::::*†Hadrosauromorpha (hadrosaurids and their closest relatives) :::::::::::::*†Hadrosauridae ("duck-billed dinosaurs"; often with crests) ::::::::::::::*†Saurolophinae (hadrosaurids with solid, small, no crests) :::::::::::::::*†Brachylophosaurini (short-crested) :::::::::::::::*†Kritosaurini (enlarged, solid nasal crests) :::::::::::::::*†Saurolophini (small, spike-like crests) :::::::::::::::*†Edmontosaurini (flat-headed saurolophines) ::::::::::::::*†Lambeosaurinae (hadrosaurids often with hollow crests) :::::::::::::::*†Aralosaurini (solid-crested) :::::::::::::::*†Tsintaosaurini (vertical, tube-like crests) :::::::::::::::*†Parasaurolophini (long, backwards-arcing crests) :::::::::::::::*†Lambeosaurini (usually rounded crests) :*
Saurischia Saurischia ( , meaning "reptile-hipped" from the Greek ' () meaning 'lizard' and ' () meaning 'hip joint') is one of the two basic divisions of dinosaurs (the other being Ornithischia), classified by their hip structure. Saurischia and Ornithis ...
::*†Herrerasauridae (early bipedal carnivores) ::*†Sauropodomorpha (herbivores with small heads, long necks, and long tails) :::*†Unaysauridae (primitive, strictly bipedal "prosauropods") :::*†Plateosauria (diverse; bipeds and quadrupeds) ::::*†Massospondylidae (long-necked, primitive sauropodomorphs) ::::*†Riojasauridae (large, primitive sauropodomorphs) :::::*†Sauropodiformes (heavy, bipeds and quadrupeds) ::::::*†Sauropoda (very large and heavy; quadrupedal) :::::::*†Lessemsauridae (gigantic yet lacking several weight-saving adaptations) :::::::*†Gravisauria ("heavy lizards") ::::::::*†Eusauropoda ("true sauropods") :::::::::*†Turiasauria (often large, widespread sauropods) :::::::::*†Neosauropoda ("new sauropods"; columnar limbs) ::::::::::*†Diplodocoidea (skulls and tails elongated; teeth typically narrow and pencil-like) :::::::::::*†Rebbachisauridae (short-necked, low-browsing diplodocoids often with high backs) :::::::::::*†Flagellicaudata (whip-tailed) ::::::::::::*†Dicraeosauridae (small, short-necked diplodocoids with enlarged cervical and dorsal vertebrae) ::::::::::::*†Diplodocidae (extremely long-necked) :::::::::::::*†Apatosaurinae (robust cervical vertebrae) :::::::::::::*†Diplodocinae (long, thin necks) ::::::::::*†Macronaria (boxy skulls; spoon- or pencil-shaped teeth) :::::::::::*†Titanosauriformes ("titan lizard forms") ::::::::::::*†Brachiosauridae (long-necked, long-armed macronarians) ::::::::::::*†Somphospondyli ("porous vertebrae") :::::::::::::*†Euhelopodidae (stocky, mostly Asian) :::::::::::::*†Titanosauria (diverse; stocky, with wide hips; most common in the Late Cretaceous of southern continents) ::*Theropoda (carnivorous) :::*†Neotheropoda ("new theropods") ::::*†Coelophysoidea (early theropods; includes ''Coelophysis'' and close relatives) ::::*†"Dilophosaur-grade neotheropods" (larger kink-snouted dinosaurs) ::::*Averostra ("bird snouts") :::::*†Ceratosauria (generally elaborately horned carnivores that existed from the Jurassic to Cretaceous periods, originally included Coelophysoidea) ::::::*†Ceratosauridae (ceratosaurs with large teeth) ::::::*†Abelisauroidea (ceratosaurs exemplified by reduced arms and hands) :::::::*†Abelisauridae (large abelisauroids with short arms and oftentimes elaborate facial ornamentation) :::::::*†Noasauridae (diverse, generally light theropods; may include several obscure taxa) ::::::::*†Elaphrosaurinae (bird-like; omnivorous as juveniles but herbivorous as adults) ::::::::*†Noasaurinae (small carnivores) :::::*Tetanurae (stiff-tailed dinosaurs) ::::::*†Megalosauroidea (early group of large carnivores) :::::::*†Piatnitzkysauridae (small basal megalosauroids endemic to the Americas) :::::::*†Megalosauridae (large megalosauroids with powerful arms and hands) :::::::*†Spinosauridae (crocodile-like, semiaquatic carnivores) ::::::*†Avetheropoda ("bird theropods") :::::::*†Megaraptora (theropods with large hand claws; either carnosaurs or coelurosaurs, potentially tyrannosauroids) :::::::*†Carnosauria (large meat-eating dinosaurs; megalosauroids sometimes included) ::::::::*†Metriacanthosauridae (primitive Asian allosauroids) ::::::::*†Allosauridae (''Allosaurus'' and its very closest relatives) ::::::::*†Carcharodontosauria (robust allosauroids) :::::::::*†Carcharodontosauridae (includes some of the largest purely terrestrial carnivores) :::::::::*†Neovenatoridae ("new hunters"; may include megaraptorans) :::::::*Coelurosauria (feathered theropods, with a range of body sizes and niches) ::::::::*†"Nexus of basal coelurosaurs" (used by Tweet to denote well-known taxa with unstable positions at the base of Coelurosauria) ::::::::*†Tyrannoraptora ("tyrant thieves") :::::::::*†Compsognathidae (small early coelurosaurs with short forelimbs) :::::::::*†Tyrannosauroidea (mostly large, primitive coelurosaurs) ::::::::::*†Proceratosauridae (tyrannosauroids with head crests) ::::::::::*†Tyrannosauridae (''Tyrannosaurus'' and close relatives) :::::::::*†Maniraptoriformes (bird-like dinosaurs) ::::::::::*†Ornithomimosauria (small-headed, mostly toothless, omnivorous or possible herbivores) :::::::::::*†Ornithomimidae (very ostrich-like dinosaurs) ::::::::::*
Maniraptora Maniraptora is a clade of coelurosaurian dinosaurs which includes the birds and the non-avian dinosaurs that were more closely related to them than to ''Ornithomimus velox''. It contains the major subgroups Avialae, Deinonychosauria, Oviraptoros ...
(dinosaurs with pennaceous feathers) :::::::::::*†Alvarezsauroidea (small hunters with reduced forelimbs) ::::::::::::*†Alvarezsauridae (insectivores with only one enlarged digit) :::::::::::*†Therizinosauria (tall, long-necked theropods; omnivores and herbivores) ::::::::::::*†Therizinosauroidea (larger therizinosaurs) :::::::::::::*†Therizinosauridae (sloth-like herbivores, often with enlarged claws) :::::::::::*†Oviraptorosauria (omnivorous, beaked dinosaurs) ::::::::::::*†Caudipteridae (bird-like, basal oviraptorosaurs) ::::::::::::*†Caenagnathoidea (cassowary-like oviraptorosaurs) :::::::::::::*†Caenagnathidae (toothless oviraptorosaurs known from North America and Asia) :::::::::::::*†Oviraptoridae (characterized by two bony projections at the back of the mouth; exclusive to Asia) :::::::::::*Paraves (avialans and their closest relatives) ::::::::::::*†Scansoriopterygidae (small tree-climbing theropods with membranous wings) ::::::::::::*†Deinonychosauria (toe-clawed dinosaurs; may not form a natural group) :::::::::::::*†Archaeopterygidae (small, winged theropods or primitive birds) :::::::::::::*†Troodontidae (omnivores; enlarged brain cavities) :::::::::::::*†Dromaeosauridae ("raptors") ::::::::::::::*†Microraptoria (characterized by large wings on both the arms and legs; may have been capable of powered flight) ::::::::::::::*†Eudromaeosauria (hunters with greatly enlarged sickle claws) :::::::::::::*†Unenlagiidae (piscivores; may be dromaeosaurids) ::::::::::::::*†Halszkaraptorinae (duck-like; potentially semiaquatic) ::::::::::::::*†Unenlagiinae (long-snouted) ::::::::::::*Avialae (modern birds and extinct relatives)


Timeline of major groups

Timeline of major dinosaur groups per Holtz (2007). ImageSize = width:1000px height:auto barincrement:15px PlotArea = left:10px bottom:50px top:10px right:10px Period = from:-251 till:0 TimeAxis = orientation:horizontal ScaleMajor = unit:year increment:10 start:-250 ScaleMinor = unit:year increment:1 start:-251 TimeAxis = orientation:hor AlignBars = justify Legend = orientation:vertical position:bottom columns:1 Colors = #legends id:CAR value:claret id:ANK value:rgb(0.4,0.3,0.196) id:HER value:teal id:HAD value:green id:OMN value:blue id:black value:black id:white value:white id:mesozoic value:rgb(0.54,0.54,0.258) id:triassic value:rgb(0.51,0.17,0.57) id:earlytriassic value:rgb(0.6,0.22,0.61) id:middletriassic value:rgb(0.73,0.53,0.71) id:latetriassic value:rgb(0.78,0.65,0.8) id:jurassic value:rgb(0.2,0.7,0.79) id:earlyjurassic value:rgb(0,0.69,0.89) id:middlejurassic value:rgb(0.52,0.81,0.91) id:latejurassic value:rgb(0.74,0.89,0.97) id:cretaceous value:rgb(0.5,0.78,0.31) id:earlycretaceous value:rgb(0.63,0.78,0.65) id:latecretaceous value:rgb(0.74,0.82,0.37) id:cenozoic value:rgb(0.54,0.54,0.258) id:paleogene value:rgb(0.99,0.6,0.32) id:paleocene value:rgb(0.99,0.65,0.37) id:eocene value:rgb(0.99,0.71,0.42) id:oligocene value:rgb(0.99,0.75,0.48) id:neogene value:rgb(0.999999,0.9,0.1) id:miocene value:rgb(0.999999,0.999999,0) id:pliocene value:rgb(0.97,0.98,0.68) id:quaternary value:rgb(0.98,0.98,0.5) id:pleistocene value:rgb(0.999999,0.95,0.68) id:holocene value:rgb(0.999,0.95,0.88) id:her value:red Legend:Herrerasauria id:pur value:purple Legend:Sauropodomorpha id:ther value:orange Legend:Theropoda id:orn value:green Legend:Ornithischia Legend = columns:1 left:100 top:20 columnwidth:100 BarData= bar:eratop bar:space bar:periodtop bar:space bar:NAM1 bar:NAM2 bar:NAM3 bar:NAM4 bar:NAM5 bar:NAM6 bar:NAM7 bar:NAM8 bar:NAM9 bar:NAM10 bar:NAM11 bar:NAM12 bar:NAM13 bar:NAM14 bar:NAM15 bar:NAM16 bar:NAM17 bar:NAM18 bar:NAM19 bar:NAM20 bar:NAM21 bar:NAM22 bar:NAM23 bar:NAM24 bar:NAM25 bar:NAM26 bar:NAM27 bar:NAM28 bar:NAM29 bar:NAM30 bar:space bar:period bar:space bar:era PlotData= align:center textcolor:black fontsize:M mark:(line,black) width:25 shift:(2,-5) bar:periodtop from: -251 till: -245 color:earlytriassic text:Early Triassic, Ear. from: -245 till: -228 color:middletriassic text:Middle Triassic, Middle from: -228 till: -199.6 color:latetriassic text:Late Triassic, Late from: -199.6 till: -175.6 color:earlyjurassic text:Early Jurassic, Early from: -175.6 till: -161.2 color:middlejurassic text:Middle Jurassic, Middle from: -161.2 till: -145.5 color:latejurassic text:Late Jurassic, Late from: -145.5 till: -99.6 color:earlycretaceous text:Early Cretaceous, Early from: -99.6 till: -65.5 color:latecretaceous text:Late Cretaceous, Late from: -65.5 till: -55.8 color:paleocene text:Paleocene, Paleo. from: -55.8 till: -33.9 color:eocene text:Eocene, Eocene from: -33.9 till: -23.03 color:oligocene text:Oligocene, Oligo. from: -23.03 till: -5.332 color:miocene text:Miocene, Mio. from: -5.332 till: -2.588 color:pliocene text:Pliocene, Pl. from: -2.588 till: -0.0117 color:pleistocene text:Pleistocene, Pl. from: -0.0117 till: 0 color:holocene text:Holocene, H. bar:eratop from: -251 till: -199.6 color:triassic text:
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
from: -199.6 till: -145.5 color:jurassic text:
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
from: -145.5 till: -65.5 color:cretaceous text:
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
from: -65.5 till: -23.03 color:paleogene text:Paleogene from: -23.03 till: -2.588 color:neogene text:Neogene from: -2.588 till: 0 color:quaternary text:Quaternary, Q. PlotData= align:left fontsize:M mark:(line,white) width:5 anchor:till shift:(5,-4) color:her bar:NAM1 from:-233.23 till:-210 text:Herrerasauridae color:pur bar:NAM2 from:-231.4 till:-208 text:Guaibasauridae color:pur bar:NAM3 from:-225 till:-190 text:Plateosauridae color:pur bar:NAM4 from:-228 till:-213 text:Riojasauridae color:pur bar:NAM5 from:-227 till:-176 text:Massospondyildae color:pur bar:NAM6 from:-183 till:-175 text:Vulcanodontidae color:pur bar:NAM7 from:-168 till:-125 text:Turiasauria color:pur bar:NAM8 from:-175 till:-150 text:Cetiosauridae color:pur bar:NAM9 from:-174 till:-93 text:Diplodocoidea color:pur bar:NAM10 from:-157 till:-93 text:Brachiosauridae color:pur bar:NAM11 from:-140 till:-66 text:Titanosauria color:ther bar:NAM12 from:-221 till:-183 text:Coelophysoidea color:ther bar:NAM13 from:-199.3 till:-66 text:Ceratosauria color:ther bar:NAM14 from:-170 till:-85 text:Megalosauroidea color:ther bar:NAM15 from:-175.6 till:-88 text:Carnosauria color:ther bar:NAM16 from:-130 till:-66 text:Megaraptora color:ther bar:NAM17 from:-166 till:-66 text:Tyrannosauroidea color:ther bar:NAM18 from:-151.5 till:-108 text:Compsognathidae color:ther bar:NAM19 from:-140 till:-66 text:Ornithomimosauria color:ther bar:NAM20 from:-160 till:-66 text:Alvarezsauria color:ther bar:NAM21 from:-139 till:-66 text:Therizinosauria color:ther bar:NAM22 from:-130 till:-66 text:Oviraptorosauria color:ther bar:NAM23 from:-167 till:-66 text:Deinonychosauria color:ther bar:NAM24 from:-155 till:0 shift:(-45,5) text:Avialae color:orn bar:NAM25 from:-200 till:-140 text:Heterodontosauridae color:orn bar:NAM26 from:-169 till:-100 text:Stegosauria color:orn bar:NAM27 from:-170.3 till:-66 text:Ankylosauria color:orn bar:NAM28 from:-92 till:-66 text:Pachycephalosauria color:orn bar:NAM29 from:-161 till:-66 text:Ceratopsia color:orn bar:NAM30 from:-164 till:-66 text:Ornithopoda PlotData= align:center textcolor:black fontsize:M mark:(line,black) width:25 shift:(2,-5) bar:period from: -251 till: -245 color:earlytriassic text:Early Triassic, Ear. from: -245 till: -228 color:middletriassic text:Middle Triassic, Middle from: -228 till: -199.6 color:latetriassic text:Late Triassic, Late from: -199.6 till: -175.6 color:earlyjurassic text:Early Jurassic, Early from: -175.6 till: -161.2 color:middlejurassic text:Middle Jurassic, Middle from: -161.2 till: -145.5 color:latejurassic text:Late Jurassic, Late from: -145.5 till: -99.6 color:earlycretaceous text:Early Cretaceous, Early from: -99.6 till: -65.5 color:latecretaceous text:Late Cretaceous, Late from: -65.5 till: -55.8 color:paleocene text:Paleocene, Paleo. from: -55.8 till: -33.9 color:eocene text:Eocene, Eocene from: -33.9 till: -23.03 color:oligocene text:Oligocene, Oligo. from: -23.03 till: -5.332 color:miocene text:Miocene, Mio. from: -5.332 till: -2.588 color:pliocene text:Pliocene, Pl. from: -2.588 till: -0.0117 color:pleistocene text:Pleistocene, Pl. from: -0.0117 till: 0 color:holocene text:Holocene, H. bar:era from: -251 till: -199.6 color:triassic text:
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
from: -199.6 till: -145.5 color:jurassic text:
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
from: -145.5 till: -65.5 color:cretaceous text:
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
from: -65.5 till: -23.03 color:paleogene text:Paleogene from: -23.03 till: -2.588 color:neogene text:Neogene from: -2.588 till: 0 color:quaternary text:Quaternary, Q.


Paleobiology

Knowledge about dinosaurs is derived from a variety of fossil and non-fossil records, including fossilized bones, feces, Fossil trackway, trackways, gastroliths, feathers, impressions of skin, Organ (anatomy), internal organs and other soft tissues. 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). Two topics of particular interest and study have been dinosaur size and behavior.


Size

Current evidence suggests that dinosaur average size varied through the Triassic, Early Jurassic, Late Jurassic and Cretaceous. Predatory theropod dinosaurs, which occupied most terrestrial carnivore niches during the Mesozoic, most often fall into the category when sorted by estimated weight into categories based on order of magnitude, whereas Holocene, recent predatory carnivoran mammals peak in the category. The Mode (statistics), mode of Mesozoic dinosaur body masses is between . This contrasts sharply with the average size of Cenozoic mammals, estimated by the National Museum of Natural History as about . 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. 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.


Largest and smallest

Scientists will probably never be certain of the largest organisms, 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. The tallest and heaviest dinosaur known from good skeletons is ''Giraffatitan, 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 Natural History Museum, Berlin, Museum für Naturkunde in Berlin; this mount is tall and long, and would have belonged to an animal that weighed between and  kilograms ( and  lb). The longest complete dinosaur is the long ''Diplodocus'', which was discovered in Wyoming in the United States and displayed in Pittsburgh's Carnegie Museum of Natural History in 1907. The longest dinosaur known from good fossil material is the ''Patagotitan'': the skeleton mount in the American Museum of Natural History in New York City, New York is long. The Museo Carmen Funes, Museo Municipal Carmen Funes in Plaza Huincul, Argentina, has an ''Argentinosaurus'' reconstructed skeleton mount that is long. 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 to  kilograms (90 to 110 short tons) and reached lengths of ; some of the longest were the long ''Diplodocus hallorum'' (formerly ''Seismosaurus''), the long ''Supersaurus'', and long ''Patagotitan''; and the tallest, the 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 Vertebra#Structure, neural arch described in 1878. Extrapolating from the illustration of this bone, the animal may have been long and weighed kg ( lb). 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. The largest carnivorous dinosaur was ''Spinosaurus'', reaching a length of , and weighing . Other large carnivorous theropods included ''Giganotosaurus'', ''Carcharodontosaurus'' and ''Tyrannosaurus''. ''Therizinosaurus'' and ''Deinocheirus'' were among the tallest of the theropods. The largest ornithischian dinosaur was probably the hadrosaurid ''Shantungosaurus, Shantungosaurus giganteus'' which measured . The largest individuals may have weighed as much as . The smallest dinosaur known is the bee hummingbird, with a length of only and mass of around . The smallest known non-Avialae, avialan dinosaurs were about the size of pigeons and were those theropods most closely related to birds. For example, ''Anchiornis huxleyi'' is currently the smallest non-avialan dinosaur described from an adult specimen, with an estimated weight of and a total skeletal length of . The smallest herbivorous non-avialan dinosaurs included ''Microceratus'' and ''Wannanosaurus'', at about long each.


Behavior

Many modern birds are highly social, often found living in flocks. There is general agreement that some behaviors that are common in birds, as well as in crocodiles (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 (ecology), habitat, computer simulations of their biomechanics, and comparisons with modern animals in similar ecological niches. The first potential evidence for herding or flocking (behavior), 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. 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, although there is no evidence for specific herd structures. Congregating into herds may have evolved for defense, for Bird migration, 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. The interpretation of dinosaurs as gregarious has also extended to depicting carnivorous theropods as pack hunters working together to bring down large prey. However, this lifestyle is uncommon among modern birds, crocodiles, and other reptiles, and the taphonomy, 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. The crests and frills of some dinosaurs, like the marginocephalians, theropods and Lambeosaurinae, 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 territory (animal), territorialism. Head wounds from bites suggest that theropods, at least, engaged in active aggressive confrontations. 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'', providing evidence that dinosaurs did indeed attack each other. 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. Cannibalism (zoology), Cannibalism amongst some species of dinosaurs was confirmed by tooth marks found in Madagascar in 2003, involving the theropod ''Majungasaurus''. Comparisons between the sclerotic ring, 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 Diurnality, diurnal. Based on fossil evidence from dinosaurs such as ''Oryctodromeus'', some ornithischian species seem to have led a partially fossorial (burrowing) lifestyle. Many modern birds are arboreal (tree climbing), and this was also true of many Mesozoic birds, especially the enantiornithines. While some early bird-like species may have already been arboreal as well (including dromaeosaurids) such as ''Microraptor'') 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, whether diplodocids could create sonic booms via whip-like tail snapping, and whether sauropods could float.


Communication

Modern birds are known to Animal communication, 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. Reconstruction of the plumage color of ''Anchiornis'', suggest the importance of color in visual communication in non-avian dinosaurs. 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 (bird anatomy), syrinx, suggesting they evolved independently and their common ancestor was mute. The earliest remains of a syrinx, which has enough mineral content for fossilization, was found in a specimen of the duck-like ''Vegavis, 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. 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. The crests of the Lambeosaurini and nasal chambers of ankylosaurids have been suggested to have functioned in vocal resonance, 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.


Reproductive biology

All dinosaurs laid Amniote, 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. Some species of modern bird have no nests; the cliff-nesting Common murre, 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 Broodiness#Broodiness in non-avian animals, brooding until all eggs were laid. When laying eggs, females grow a special type of bone between the hard outer bone and the Bone marrow, 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. 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 (paleontologist), 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. A specimen of the oviraptoridae, oviraptorid ''Citipati, Citipati osmolskae'' was discovered in a Chicken#Broodiness, chicken-like brooding position in 1993, which may indicate that they had begun using an insulating layer of feathers to keep the eggs warm. An embryo of the basal sauropodomorph ''Massospondylus'' was found without teeth, indicating that some parental care was required to feed the young dinosaurs. Trackways have also confirmed parental behavior among ornithopods from the Isle of Skye in northwestern Scotland. However, there is ample evidence of precociality or Precociality#Superprecociality, superprecociality among many dinosaur species, particularly theropods. For instance, non-Euornithes, 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. Both ''Tyrannosaurus'' and ''Troodon'' had juveniles with clear superprecociality and likely occupying different ecological niches than the adults. Superprecociality has been inferred for sauropods. 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.


Physiology

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. 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. 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, which is termed mesothermy by some. 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. 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. One of the greatest contributions to the modern understanding of dinosaur physiology has been histology, paleohistology, the study of microscopic tissue structure in dinosaurs. 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, though not universally present. This has led to a significant body of work in reconstructing growth curve (biology), growth curves and modeling the evolution of growth rates across various dinosaur lineages,For examples of this work conducted on different dinosaur lineages, see * * * * * * which has suggested overall that dinosaurs grew faster than living reptiles. Other lines of evidence suggesting endothermy include the presence of feathers and other types of body coverings in many lineages (see ); 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 (although these ratios can be altered during fossilization); and the discovery of South Polar region of the Cretaceous#Dinosaurs, polar dinosaurs, which lived in Australia, Antarctica, and Alaska when these places would have had cool, temperate climates. 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. Such respiratory systems, which may have appeared in the earliest saurischians, 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. The rapid airflow would also have been an effective cooling mechanism, which in conjunction with a lower metabolic rate would have prevented large sauropods from overheating. These traits may have enabled sauropods to grow quickly to gigantic sizes. 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. 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. In most living species, uric acid is excreted along with feces as a semisolid waste. However, at least some modern birds (such as hummingbirds) can be facultatively ammonotelic, excreting most of the nitrogenous wastes as ammonia. This material, as well as the output of the intestines, emerges from the cloaca. In addition, many species regurgitate Pellet (ornithology), pellets, and fossil pellets are known as early as the Jurassic from ''Anchiornis''. 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. 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''.


Origin of birds

The possibility that dinosaurs were the ancestors of birds was first suggested in 1868 by Thomas Henry Huxley. After the work of Gerhard Heilmann in the early 20th century, the Scientific theory, theory of birds as dinosaur descendants was abandoned in favor of the idea of them being descendants of generalized Thecodontia, thecodonts, with the key piece of evidence being the supposed lack of clavicles in dinosaurs. However, as later discoveries showed, clavicles (or a single fused furcula, wishbone, which derived from separate clavicles) were not actually absent; they had been found as early as 1924 in ''Oviraptor'', but misidentified as an interclavicle. In the 1970s, Ostrom revived the dinosaur–bird theory, which gained momentum in the coming decades with the advent of cladistic analysis, and a great increase in the discovery of small theropods and early birds. 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. Birds share over a hundred distinct anatomical features with theropod dinosaurs, which are now generally accepted to have been their closest ancient relatives. They are most closely allied with maniraptoran coelurosaurs. 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, or that maniraptoran theropods are the ancestors of birds but themselves are not dinosaurs, only convergent evolution, convergent with dinosaurs.


Feathers

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. Direct fossil evidence of feathers or feather-like structures has been discovered in a diverse array of species in many non-avian dinosaur groups, both among saurischians and ornithischians. Simple, branched, feather-like structures are known from Heterodontosauridae, heterodontosaurids, primitive neornithischians, and theropods, 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. Feather-like structures known as Pterosaur#Pycnofibers, pycnofibres have also been found in pterosaurs, suggesting the possibility that feather-like filaments may have been common in the bird lineage and evolved before the appearance of dinosaurs themselves. 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. ''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''. 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 Lagerstätten, 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. 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, and that maniraptoran dinosaurs with vaned feathers were not actually dinosaurs, but convergent with dinosaurs. 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.


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 Shoulder girdle, pectoral girdle, furcula (wishbone), and Keel (bird anatomy), breast bone. Comparison of bird and dinosaur skeletons through cladistic analysis strengthens the case for the link.


Soft anatomy

Large meat-eating dinosaurs had a complex system of air sacs similar to those found in modern birds, according to a 2005 investigation led by Patrick M. O'Connor. The lungs of theropod dinosaurs (carnivores that walked on two legs and had bird-like feet) likely pumped air into hollow sacs in their skeletons, as is the case in birds. "What was once formally considered unique to birds was present in some form in the ancestors of birds", O'Connor said. In 2008, scientists described ''Aerosteon, Aerosteon riocoloradensis'', the skeleton of which supplies the strongest evidence to date of a dinosaur with a bird-like breathing system. CT scanning of ''Aerosteons fossil bones revealed evidence for the existence of air sacs within the animal's body cavity.


Behavioral evidence

Fossils of the troodonts ''Mei long, Mei'' and ''Sinornithoides'' demonstrate that some dinosaurs slept with their heads tucked under their arms. This behavior, which may have helped to keep the head warm, is also characteristic of modern birds. Several Deinonychosauria, deinonychosaur and oviraptorosaur specimens have also been found preserved on top of their nests, likely brooding in a bird-like manner. 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. 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.


Extinction of major groups

All non-avian dinosaurs and most lineages of birds became extinct in a extinction event, mass extinction event, called the Cretaceous–Paleogene extinction event, 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, 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. Alongside dinosaurs, many other groups of animals became extinct: pterosaurs, marine reptiles such as mosasaurs and plesiosaurs, several groups of mammals, ammonoidea, ammonites (nautilus-like Mollusca, mollusks), rudists (reef-building bivalvia, bivalves), and various groups of marine plankton. In all, approximately 47% of genera and 76% of species on Earth became extinct during the K-Pg extinction event. 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; 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 seed predation, eating seeds for survival.


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. 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. However, these comparisons did not account either for varying preservation potential between rock units or for different extents of exploration and excavation. In 1984, Dale Russell carried out an analysis to account for these biases, and found no evidence of a decline; another analysis by David Fastovsky and colleagues in 2004 even showed that dinosaur diversity continually increased until the extinction, but this analysis has been rebutted. Since then, different approaches based on statistics and mathematical models have variously supported either a sudden extinction or a gradual decline. 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.


Impact event

The Alvarez hypothesis, bolide impact hypothesis, first brought to wide attention in 1980 by Walter Alvarez, Luis Walter Alvarez, Luis Alvarez, and colleagues, attributes the K-Pg extinction event to a bolide (extraterrestrial projectile) impact. 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. Shocked quartz, indicative of a strong shockwave emanating from an impact, was also found worldwide. The actual impact site remained elusive until a Chicxulub crater, crater measuring wide was discovered in the Yucatán Peninsula of southeastern Mexico, and was publicized in a 1991 paper by Alan R. Hildebrand, Alan Hildebrand and colleagues. Now, the bulk of the evidence suggests that a bolide wide impacted the Yucatán Peninsula 66 million years ago, forming this crater and creating a "kill mechanism" that triggered the extinction event. Within hours, the Chicxulub impact would have created immediate effects such as earthquakes, tsunamis, and a global firestorm that likely killed unsheltered animals and started wildfires. 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. 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. (This role was ascribed to sulphate aerosols until experiments demonstrated otherwise.) 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.


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. Greenhouse gases such as carbon dioxide and sulphur dioxide would have been released by this volcanic activity, resulting in climate change through temperature perturbations of roughly but possibly as high as . Like the Chicxulub impact, the eruptions may also have released sulphate aerosols, which would have caused acid rain and global cooling. 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. 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; even after the crater's discovery, the impact was still thought to only have had a regional, not global, effect on the extinction event. In response, Luis Alvarez rejected volcanic activity as an explanation for the iridium layer and the extinction as a whole. 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. Some estimates have placed the start of the second phase in the Deccan Traps eruptions within 50,000 years after the Chicxulub impact. 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. 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. They consider the Chicxulub impact to have worsened the ongoing climate change caused by the eruptions. 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. 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.


Possible Paleocene survivors

Non-avian dinosaur remains have occasionally been found above the K-Pg boundary. In 2000, Spencer G. Lucas, 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. If the bone was not deposition (geology), 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. Other evidence includes the presence of dinosaur remains in the Hell Creek Formation up to 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. However, these supposed Paleocene dinosaurs are considered by many other researchers to be reworked fossil, reworked, that is, washed out of their original locations and then re-buried in younger sediments. The age estimates have also been considered unreliable.


Cultural depictions

By human standards, dinosaurs were creatures of fantastic appearance and often enormous size. As such, they have captured the popular imagination and become an enduring part of human culture. 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. Public enthusiasm for dinosaurs first developed in Victorian era, Victorian England, where in 1854, three decades after the first scientific descriptions of dinosaur remains, a menagerie of lifelike Crystal Palace dinosaurs, 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 Natural history museum, museums around the world, ensuring that successive generations would be introduced to the animals in an immersive and exciting way. 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. The popular preoccupation with dinosaurs has ensured their appearance in literature, film, and other Media (communication), media. Beginning in 1852 with a passing mention in Charles Dickens ''Bleak House'', dinosaurs have been featured in large numbers of fictional works. Jules Verne's 1864 novel ''Journey to the Center of the Earth'', Arthur Conan Doyle, Sir Arthur Conan Doyle's 1912 book ''The Lost World (Doyle novel), The Lost World'', the 1914 animated film ''Gertie the Dinosaur'' (featuring the first animated dinosaur), the iconic 1933 motion picture, film ''King Kong (1933 film), King Kong'', the 1954 ''Godzilla (1954 film), Godzilla'' and its many sequels, the best-selling 1990 novel ''Jurassic Park (novel), Jurassic Park'' by Michael Crichton and its 1993 Jurassic Park (film), 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 Company (law), 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.


See also

* Dinosaur diet and feeding * Evolutionary history of life * Lists of dinosaur-bearing stratigraphic units * List of dinosaur genera * List of bird genera * List of birds * List of informally named dinosaurs


Further reading

* * * . * *


Notes


Bibliography

* * * * * * * The 5th edition of the book is available from th
Internet Archive
Retrieved 2019-10-19. * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * "Reprint of papers published in a special volume of Modern geology [v. 18 (Halstead memorial volume), 1993], with five additional contributions.--Pref." * *


References

{{DEFAULTSORT:Dinosaur Carnian first appearances Dinosaurs, Extant Late Triassic first appearances Fossil taxa described in 1842 Taxa named by Richard Owen