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Dinosaur

Dinosaur

Overview
Dinosaurs are a diverse group of animals of the clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...

 and superorder Dinosauria. They were the dominant terrestrial vertebrate
Vertebrate
Vertebrates are animals that are members of the subphylum Vertebrata . Vertebrates are the largest group of chordates, with currently about 58,000 species described. Vertebrates include the jawless fishes, bony fishes, sharks and rays, amphibians, reptiles, mammals, and birds...

s for over 160 million years, from the late Triassic
Triassic
The Triassic is a geologic period and system that extends from about 250 to 200 Mya . As the first period of the Mesozoic Era, the Triassic follows the Permian and is followed by the Jurassic. Both the start and end of the Triassic are marked by major extinction events...

 period (about 230 million years ago) until the end of the Cretaceous
Cretaceous
The Cretaceous , derived from the Latin "creta" , usually abbreviated K for its German translation Kreide , is a geologic period and system from circa to million years ago. In the geologic timescale, the Cretaceous follows the Jurassic period and is followed by the Paleogene period of the...

 (about 65 million years ago), when the Cretaceous–Paleogene extinction event led to the extinction of all non-avian dinosaurs at the close of the Mesozoic era.
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Encyclopedia
Dinosaurs are a diverse group of animals of the clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...

 and superorder Dinosauria. They were the dominant terrestrial vertebrate
Vertebrate
Vertebrates are animals that are members of the subphylum Vertebrata . Vertebrates are the largest group of chordates, with currently about 58,000 species described. Vertebrates include the jawless fishes, bony fishes, sharks and rays, amphibians, reptiles, mammals, and birds...

s for over 160 million years, from the late Triassic
Triassic
The Triassic is a geologic period and system that extends from about 250 to 200 Mya . As the first period of the Mesozoic Era, the Triassic follows the Permian and is followed by the Jurassic. Both the start and end of the Triassic are marked by major extinction events...

 period (about 230 million years ago) until the end of the Cretaceous
Cretaceous
The Cretaceous , derived from the Latin "creta" , usually abbreviated K for its German translation Kreide , is a geologic period and system from circa to million years ago. In the geologic timescale, the Cretaceous follows the Jurassic period and is followed by the Paleogene period of the...

 (about 65 million years ago), when the Cretaceous–Paleogene extinction event led to the extinction of all non-avian dinosaurs at the close of the Mesozoic era. The fossil
Fossil
Fossils are the preserved remains or traces of animals , plants, and other organisms from the remote past...

 record indicates that bird
Bird
Birds are feathered, winged, bipedal, endothermic , egg-laying, vertebrate animals. Around 10,000 living species and 188 families makes them the most speciose class of tetrapod vertebrates. They inhabit ecosystems across the globe, from the Arctic to the Antarctic. Extant birds range in size from...

s evolved
Evolution
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...

 within theropod dinosaurs during the Jurassic
Jurassic
The Jurassic is a geologic period and system that extends from about Mya to  Mya, that is, from the end of the Triassic to the beginning of the Cretaceous. The Jurassic constitutes the middle period of the Mesozoic era, also known as the age of reptiles. The start of the period is marked by...

 period. Some of them survived the Cretaceous–Paleogene extinction event, including the ancestors of all modern birds. Consequently, in modern classification systems, birds are considered a type of dinosaur—the only group which survived to the present day.

Dinosaurs are a varied group of animals. Birds, at over 9,000 living species, are the most diverse group of vertebrates besides perciform
Perciformes
The Perciformes, also called the Percomorphi or Acanthopteri, is one of the largest orders of vertebrates, containing about 40% of all bony fish. Perciformes means perch-like. They belong to the class of ray-finned fish and comprise over 7,000 species found in almost all aquatic environments...

 fish. Using fossil evidence, paleontologists have identified over 500 distinct genera
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...

 and more than 1,000 different species of non-avian dinosaurs. Dinosaurs are represented on every continent by both extant species and fossil remains. Some are herbivorous, others carnivorous. Many dinosaurs have been bipedal, and many extinct groups were also quadrupedal, and some were able to shift between these body postures. Many species possess elaborate display structures such as horns or crests, and some prehistoric groups even developed skeletal modifications such as bony armor
Armour (zoology)
Armour in animals is external or superficial protection against attack by predators, formed as part of the body , usually through the hardening of body tissues, outgrowths or secretions. It has therefore mostly developed in 'prey' species...

 and spines
Spine (zoology)
A spine is a hard, thorny or needle-like structure which occurs on various animals. Animals such as porcupines and sea urchins grow spines as a self-defense mechanism. Spines are often formed of keratin...

. Avian dinosaurs have been the planet's dominant flying vertebrate since the extinction of the pterosaur
Pterosaur
Pterosaurs were flying reptiles of the clade or order Pterosauria. They existed from the late Triassic to the end of the Cretaceous Period . Pterosaurs are the earliest vertebrates known to have evolved powered flight...

s, and evidence suggests that all ancient dinosaurs built nests and laid eggs much as avian species do today. Dinosaurs varied widely in size and weight; the smallest adult theropods were less than 100 centimeters (40 inches) long, while the largest sauropods could reach lengths of almost 50 meters (165 feet) and were several stories tall.

Although the word dinosaur means "terrible lizard," the name is somewhat misleading, as dinosaurs were not lizard
Lizard
Lizards are a widespread group of squamate reptiles, with nearly 3800 species, ranging across all continents except Antarctica as well as most oceanic island chains...

s. Rather, they were a separate group of reptiles with a distinct upright posture not found in lizards. Through the first half of the 20th century, most of the scientific community believed dinosaurs were sluggish, unintelligent, and cold-blooded
Poikilotherm
A poikilotherm is an organism whose internal temperature varies considerably. It is the opposite of a homeotherm, an organism which maintains thermal homeostasis. Usually the variation is a consequence of variation in the ambient environmental temperature...

. Most research conducted since the 1970s
Dinosaur renaissance
The dinosaur renaissance was a small-scale scientific revolution that started in the late 1960s, and led to renewed academic and popular interest in dinosaurs...

, however, has indicated that dinosaurs were active animals with elevated metabolism
Metabolism
Metabolism is the set of chemical reactions that happen in the cells of living organisms to sustain life. These processes allow organisms to grow and reproduce, maintain their structures, and respond to their environments. Metabolism is usually divided into two categories...

s and numerous adaptations for social interaction, and many groups (especially the carnivores) were among the most intelligent organisms of the time.

Since the first dinosaur fossil
Fossil
Fossils are the preserved remains or traces of animals , plants, and other organisms from the remote past...

s were recognized in the early 19th century, mounted fossil dinosaur skeletons or replicas have been major attractions at museums around the world, and dinosaurs have become a part of world culture. Their diversity, the large sizes of some groups, and their seemingly monstrous and fantastic nature have captured the interest and imagination of the general public for over a century. They have been featured in best-selling books and films such as Jurassic Park
Jurassic Park (film)
Jurassic Park is a 1993 American science fiction adventure film directed by Steven Spielberg. The film is based on the novel of the same name by Michael Crichton. It stars Sam Neill, Laura Dern, Jeff Goldblum, Richard Attenborough, Martin Ferrero, and Bob Peck...

, and new discoveries are regularly covered by the media.

Etymology


The taxon
Taxon
|thumb|270px|[[African elephants]] form a widely-accepted taxon, the [[genus]] LoxodontaA taxon is a group of organisms, which a taxonomist adjudges to be a unit. Usually a taxon is given a name and a rank, although neither is a requirement...

 Dinosauria was formally named in 1842 by paleontologist Sir Richard Owen
Richard Owen
Sir Richard Owen, FRS KCB was an English biologist, comparative anatomist and palaeontologist.Owen is probably best remembered today for coining the word Dinosauria and for his outspoken opposition to Charles Darwin's theory of evolution by natural selection...

, who used it to refer to the "distinct tribe or sub-order of Saurian Reptiles" that were then being recognized in England and around the world. The term is derived from the Greek
Ancient Greek
Ancient Greek is the stage of the Greek language in the periods spanning the times c. 9th–6th centuries BC, , c. 5th–4th centuries BC , and the c. 3rd century BC – 6th century AD of ancient Greece and the ancient world; being predated in the 2nd millennium BC by Mycenaean Greek...

 words δεινός (deinos, meaning "terrible," "potent," or "fearfully great") and σαῦρος (sauros, meaning "lizard" or "reptile"). Though the taxonomic name has often been interpreted as a reference to dinosaurs' teeth, claws, and other fearsome characteristics, Owen intended it merely to evoke their size and majesty.

Modern definition



Formal definitions are written to correspond with scientific conceptions of dinosaurs that predate the modern use of phylogenetics
Phylogenetics
In biology, phylogenetics is the study of evolutionary relatedness among groups of organisms , which is discovered through molecular sequencing data and morphological data matrices...

. The continuity of meaning is intended to prevent confusion about what the term "dinosaur" means.

Under phylogenetic taxonomy, dinosaurs are usually defined as the group consisting of "Triceratops
Triceratops
Triceratops is a genus of herbivorous ceratopsid dinosaur which lived during the late Maastrichtian stage of the Late Cretaceous Period, around 68 to 65 million years ago in what is now North America. It was one of the last dinosaur genera to appear before the great Cretaceous–Paleogene...

, Neornithes [modern birds], their most recent common ancestor
Most recent common ancestor
In genetics, the most recent common ancestor of any set of organisms is the most recent individual from which all organisms in the group are directly descended...

, and all descendants". It has also been suggested that Dinosauria be defined with respect to the MRCA
Most recent common ancestor
In genetics, the most recent common ancestor of any set of organisms is the most recent individual from which all organisms in the group are directly descended...

 of Megalosaurus
Megalosaurus
Megalosaurus is a genus of large meat-eating theropod dinosaurs of the Middle Jurassic period of Europe...

and Iguanodon
Iguanodon
Iguanodon is a genus of ornithopod dinosaur that lived roughly halfway between the first of the swift bipedal hypsilophodontids and the ornithopods' culmination in the duck-billed dinosaurs...

, because these were two of the three genera cited by Richard Owen when he recognized the Dinosauria. Both definitions result in the same set of animals being defined as dinosaurs, that is "Dinosauria = Ornithischia
Ornithischia
Ornithischia or Predentata is an extinct order of beaked, herbivorous dinosaurs. The name ornithischia is derived from the Greek ornitheos meaning 'of a bird' and ischion meaning 'hip joint'...

 + Saurischia
Saurischia
Saurischia meaning 'lizard' and ischion meaning 'hip joint') is one of the two orders, or basic divisions, of dinosaurs. In 1888, Harry Seeley classified dinosaurs into two orders, based on their hip structure...

", which encompasses theropods
Theropoda
Theropoda is both a suborder of bipedal saurischian dinosaurs, and a clade consisting of that suborder and its descendants . Dinosaurs belonging to the suborder theropoda were primarily carnivorous, although a number of theropod groups evolved herbivory, omnivory, and insectivory...

 (mostly bipedal carnivore
Carnivore
A carnivore meaning 'meat eater' is an organism that derives its energy and nutrient requirements from a diet consisting mainly or exclusively of animal tissue, whether through predation or scavenging...

s and bird
Bird
Birds are feathered, winged, bipedal, endothermic , egg-laying, vertebrate animals. Around 10,000 living species and 188 families makes them the most speciose class of tetrapod vertebrates. They inhabit ecosystems across the globe, from the Arctic to the Antarctic. Extant birds range in size from...

s), ankylosauria
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. Ankylosaurs were bulky quadrupeds, with short, powerful limbs. They are first known to have appeared in the early Jurassic Period of...

ns (armored herbivorous quadrupeds), stegosauria
Stegosauria
Known colloquially as stegosaurs, the Stegosauria are a group of herbivorous dinosaurs of the Jurassic and Early Cretaceous Periods, being found mostly in the Northern Hemisphere, predominantly in what is now North America and China....

ns (plated herbivorous quadrupeds), ceratopsia
Ceratopsia
Ceratopsia or Ceratopia is a group of herbivorous, beaked dinosaurs which thrived in what are now North America, Europe, and Asia, during the Cretaceous Period, although ancestral forms lived earlier, in the Jurassic. The earliest known ceratopsian, Yinlong downsi, lived between 161.2 and 155.7...

ns (herbivorous quadrupeds with horns and frills), ornithopods (bipedal or quadrupedal herbivores including "duck-bills"), and, perhaps, sauropodomorphs
Sauropodomorpha
Sauropodomorpha is an extinct clade of long-necked, herbivorous, saurischian dinosaurs which includes the sauropods and their ancestral relatives. Sauropods generally grew to very large sizes, had long necks and tails, were quadrupedal, and became the largest animals to ever walk the Earth. The...

 (mostly large herbivorous
Herbivore
Herbivores are organisms that are anatomically and physiologically adapted to eat plant-based foods. Herbivory is a form of consumption in which an organism principally eats autotrophs such as plants, algae and photosynthesizing bacteria. More generally, organisms that feed on autotrophs in...

 quadruped
Quadruped
Quadrupedalism is a form of land animal locomotion using four limbs or legs. An animal or machine that usually moves in a quadrupedal manner is known as a quadruped, meaning "four feet"...

s with long necks and tails).

Many paleontologists note that the point at which sauropodomorphs and theropods diverged may omit sauropodomorphs from the definition for both saurischians and dinosaurs. To avoid the instability of Dinosauria, a more conservative definition of Dinosauria is defined with respect to four anchoring nodes: Triceratops horridus, Saltasaurus loricatus, and Passer domesticus, their most recent common ancestor, and all descendants. This "safer" definition can be expressed as "Dinosauria = Ornithischia + Sauropodomorpha
Sauropodomorpha
Sauropodomorpha is an extinct clade of long-necked, herbivorous, saurischian dinosaurs which includes the sauropods and their ancestral relatives. Sauropods generally grew to very large sizes, had long necks and tails, were quadrupedal, and became the largest animals to ever walk the Earth. The...

 + Theropoda
Theropoda
Theropoda is both a suborder of bipedal saurischian dinosaurs, and a clade consisting of that suborder and its descendants . Dinosaurs belonging to the suborder theropoda were primarily carnivorous, although a number of theropod groups evolved herbivory, omnivory, and insectivory...

".

There is a wide consensus among paleontologists that birds are the descendants of theropod dinosaurs. Using the strict phylogenetic nomenclatural
Phylogenetic nomenclature
Phylogenetic nomenclature or phylogenetic taxonomy is an alternative to rank-based nomenclature, applying definitions from cladistics . Its two defining features are the use of phylogenetic definitions of biological taxon names, and the lack of obligatory ranks...

 definition that all descendants of a single common ancestor must be included in a group for that group to be natural, birds would thus be dinosaurs and dinosaurs are, therefore, not extinct. Birds are classified by most paleontologists as belonging to the subgroup Maniraptora
Maniraptora
Maniraptora is a clade of coelurosaurian dinosaurs which includes the birds and the dinosaurs that were more closely related to them than to Ornithomimus velox. It contains the major subgroups Avialae, Deinonychosauria, Oviraptorosauria and Therizinosauria. Ornitholestes and the Alvarezsauroidea...

, which are coelurosaurs
Coelurosauria
Coelurosauria is the clade containing all theropod dinosaurs more closely related to birds than to carnosaurs. In the past, it was used to refer to all small theropods, although this classification has been abolished...

, which are theropods, which are saurischia
Saurischia
Saurischia meaning 'lizard' and ischion meaning 'hip joint') is one of the two orders, or basic divisions, of dinosaurs. In 1888, Harry Seeley classified dinosaurs into two orders, based on their hip structure...

ns, which are dinosaurs.

From the point of view of cladistics, birds are dinosaurs, but in ordinary speech the word "dinosaur" does not include birds. Additionally, referring to dinosaurs that are not birds as "non-avian dinosaurs" is cumbersome. For clarity, this article will use "dinosaur" as a synonym for "non-avian dinosaur". The term "non-avian dinosaur" will be used for emphasis as needed.

General description



Using one of the above definitions, dinosaurs (aside from birds) can be generally described as terrestrial archosaur
Archosaur
Archosaurs are a group of diapsid amniotes whose living representatives consist of modern birds and crocodilians. This group also includes all extinct non-avian dinosaurs, many extinct crocodilian relatives, and pterosaurs. Archosauria, the archosaur clade, is a crown group that includes the most...

ian reptile
Reptile
Reptiles are members of a class of air-breathing, ectothermic vertebrates which are characterized by laying shelled eggs , and having skin covered in scales and/or scutes. They are tetrapods, either having four limbs or being descended from four-limbed ancestors...

s with limbs held erect beneath the body, that existed from the Late Triassic
Late Triassic
The Late Triassic is in the geologic timescale the third and final of three epochs of the Triassic period. The corresponding series is known as the Upper Triassic. In the past it was sometimes called the Keuper, after a German lithostratigraphic group that has a roughly corresponding age...

 (first appearing in the Carnian
Carnian
The Carnian is the lowermost stage of the Upper Triassic series . It lasted from about 228.7 till 216.5 million years ago . The Carnian is preceded by the Ladinian and is followed by the Norian...

 faunal stage
Faunal stage
In chronostratigraphy, a stage is a succession of rock strata laid down in a single age on the geologic timescale, which usually represents millions of years of deposition. A given stage of rock and the corresponding age of time will by convention have the same name, and the same boundaries.Rock...

) to the Late Cretaceous
Late Cretaceous
The Late Cretaceous is the younger of two epochs into which the Cretaceous period is divided in the geologic timescale. Rock strata from this epoch form the Upper Cretaceous series...

 (going extinct at the end of the Maastrichtian
Maastrichtian
The Maastrichtian is, in the ICS' geologic timescale, the latest age or upper stage of the Late Cretaceous epoch or Upper Cretaceous series, the Cretaceous period or system, and of the Mesozoic era or erathem. It spanned from 70.6 ± 0.6 Ma to 65.5 ± 0.3 Ma...

). Many prehistoric animals are popularly conceived of as dinosaurs, such as ichthyosaur
Ichthyosaur
Ichthyosaurs were giant marine reptiles that resembled fish and dolphins...

s, mosasaur
Mosasaur
Mosasaurs are large extinct marine lizards. The first fossil remains were discovered in a limestone quarry at Maastricht on the Meuse in 1764...

s, plesiosaur
Plesiosaur
Plesiosauroidea is an extinct clade of carnivorous plesiosaur marine reptiles. Plesiosauroids, are known from the Jurassic and Cretaceous Periods...

s, pterosaur
Pterosaur
Pterosaurs were flying reptiles of the clade or order Pterosauria. They existed from the late Triassic to the end of the Cretaceous Period . Pterosaurs are the earliest vertebrates known to have evolved powered flight...

s, and Dimetrodon
Dimetrodon
Dimetrodon was a predatory synapsid genus that flourished during the Permian period, living between 280–265 million years ago ....

, but are not classified scientifically as dinosaurs. Marine reptiles like ichthyosaurs, mosasaurs, and plesiosaurs were neither terrestrial nor archosaurs; pterosaurs were archosaurs but not terrestrial; and Dimetrodon was a Permian
Permian
The PermianThe term "Permian" was introduced into geology in 1841 by Sir Sir R. I. Murchison, president of the Geological Society of London, who identified typical strata in extensive Russian explorations undertaken with Edouard de Verneuil; Murchison asserted in 1841 that he named his "Permian...

 animal more closely related to mammals. Dinosaurs were the dominant terrestrial vertebrates of the Mesozoic, especially the Jurassic
Jurassic
The Jurassic is a geologic period and system that extends from about Mya to  Mya, that is, from the end of the Triassic to the beginning of the Cretaceous. The Jurassic constitutes the middle period of the Mesozoic era, also known as the age of reptiles. The start of the period is marked by...

 and Cretaceous. Other groups of animals were restricted in size and niches; mammals, for example, rarely exceeded the size of a cat, and were generally rodent-sized carnivores of small prey. One notable exception is Repenomamus giganticus
Repenomamus
Repenomamus is the largest mammal known from the Cretaceous period of Manchuria, and it is the mammal for which there is the best evidence that it fed on dinosaurs. It is not possible to determine if Repenomamus actively hunted live dinosaurs or scavenged dead dinosaurs.-Paleobiology:Repenomamus...

, a triconodont weighing between 12 kilograms (26.5 lb) and 14 kilograms (30.9 lb) that is known to have eaten small dinosaurs like young Psittacosaurus
Psittacosaurus
Psittacosaurus is a genus of psittacosaurid ceratopsian dinosaur from the Early Cretaceous Period of what is now Asia, about 130 to 100 million years ago. It is notable for being the most species-rich dinosaur genus...

.

Dinosaurs were an extremely varied group of animals; according to a 2006 study, over 500 dinosaur genera have been identified with certainty so far, and the total number of genera preserved in the fossil record has been estimated at around 1850, nearly 75% of which remain to be discovered. An earlier study predicted that about 3400 dinosaur genera existed, including many which would not have been preserved in the fossil record., 1047 different species of dinosaurs have been named. Some were herbivorous, others carnivorous. Some dinosaurs were bipeds, some were quadrupeds, and others, such as Ammosaurus
Ammosaurus
Ammosaurus is a genus of sauropodomorph dinosaur from the Early and Middle Jurassic Period of North America. At 4 meters in length, it was small compared to some other members of its suborder, which included the largest animals ever to walk the Earth...

and Iguanodon
Iguanodon
Iguanodon is a genus of ornithopod dinosaur that lived roughly halfway between the first of the swift bipedal hypsilophodontids and the ornithopods' culmination in the duck-billed dinosaurs...

, could walk just as easily on two or four legs. Many had bony armor
Armour (zoology)
Armour in animals is external or superficial protection against attack by predators, formed as part of the body , usually through the hardening of body tissues, outgrowths or secretions. It has therefore mostly developed in 'prey' species...

, or cranial modifications like horns and crests. Although known for large size, many dinosaurs were human-sized or smaller. Dinosaur remains have been found on every continent on Earth, including Antarctica. No non-avian dinosaurs are known to have lived in marine habitats or in aerial habitats, although it is possible some feathered non-avian theropods were flyers. There is also evidence that some spinosaurids
Spinosauridae
Spinosauridae is a family of specialised theropod dinosaurs. Members of this family were large, bipedal predators with elongated, crocodile-like skulls, sporting conical teeth with no or only very tiny serrations. The front dentary teeth fanned out, giving the animal a characteristic look...

 had semi-aquatic habits.

Distinguishing anatomical features


While recent discoveries have made it more difficult to present a universally agreed-upon list of dinosaurs' distinguishing features, nearly all dinosaurs discovered so far share certain modifications to the ancestral archosaur
Archosaur
Archosaurs are a group of diapsid amniotes whose living representatives consist of modern birds and crocodilians. This group also includes all extinct non-avian dinosaurs, many extinct crocodilian relatives, and pterosaurs. Archosauria, the archosaur clade, is a crown group that includes the most...

ian skeleton. Although some later groups of dinosaurs featured further modified versions of these traits, they are considered typical across Dinosauria; the earliest dinosaurs had them and passed them on to all their descendants. Such common features across a taxonomic group are called synapomorphies
Synapomorphy
In cladistics, a synapomorphy or synapomorphic character is a trait that is shared by two or more taxa and their most recent common ancestor, whose ancestor in turn does not possess the trait. A synapomorphy is thus an apomorphy visible in multiple taxa, where the trait in question originates in...

.

A detailed assessment of archosaur interrelations by S. Nesbitt confirmed or found the following 12 unambiguous synapomorphies, some previously known:
  • in the skull, a supratemporal fossa (excavation) is present in front of the supratemporal fenestra
  • epipophyses present in anterior neck vertebrae (except atlas and axis)
  • apex of deltopectoral crest (a projection on which the deltopectoral
    Clavipectoral triangle
    The clavipectoral triangle is an anatomical region found in humans and other animals. It is bordered by the following structures:* Pectoralis major muscle* Deltoid muscle* Clavicle...

     muscles attach) located at or more than 30% down the length of the humerus
    Humerus
    The humerus is a long bone in the arm or forelimb that runs from the shoulder to the elbow....

     (upper arm bone)
  • radius
    Radius (bone)
    The radius is one of the two large bones of the forearm, the other being the ulna. It extends from the lateral side of the elbow to the thumb side of the wrist and runs parallel to the ulna, which exceeds it in length and size. It is a long bone, prism-shaped and slightly curved longitudinally...

     shorter than 80% of humerus length
  • fourth trochanter (projection where the caudofemoralis muscle attaches) on the femur
    Femur
    The femur , or thigh bone, is the most proximal bone of the leg in tetrapod vertebrates capable of walking or jumping, such as most land mammals, birds, many reptiles such as lizards, and amphibians such as frogs. In vertebrates with four legs such as dogs and horses, the femur is found only in...

     (thigh bone) is a sharp flange
  • fourth trochanter asymmetrical, with distal margin forming a steeper angle to the shaft
  • on the astragalus
    Talus bone
    -External links:* *...

     and calcaneum the proximal articular facet for fibula occupies less than 30% of the transverse width of the element
  • exocciptials (bones at the back of the skull) do not meet along the midline on the floor of the endocranial cavity
  • proximal articular surfaces of the ischium with the ilium
    Ilium (bone)
    The ilium is the uppermost and largest bone of the pelvis, and appears in most vertebrates including mammals and birds, but not bony fish. All reptiles have an ilium except snakes, although some snake species have a tiny bone which is considered to be an ilium.The name comes from the Latin ,...

     and the pubis
    Pubis (bone)
    In vertebrates, the pubic bone is the ventral and anterior of the three principal bones composing either half of the pelvis.It is covered by a layer of fat, which is covered by the mons pubis....

     separated by a large concave surface
  • cnemial crest on the tibia
    Tibia
    The tibia , shinbone, or shankbone is the larger and stronger of the two bones in the leg below the knee in vertebrates , and connects the knee with the ankle bones....

     (shinbone) arcs anterolaterally
  • distinct proximodistally oriented ridge present on the posterior face of the distal end of the tibia

Nesbitt found a number of further potential synapomorphies, and discounted a number of synapomorphies previously suggested. Some of these are also present in silesaurids
Silesauridae
Silesauridae is an extinct clade of dinosauriformes, a group of Triassic reptiles that included early ancestors of the dinosaurs. The silesaurids were a sister clade to the dinosaurs. The family was named in 2010 by paleontologist Max C. Langer and colleagues from Brazil and Argentina...

, 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 cenmial crest on the tibia and of an ascending process on the astragalus, and many others.



A variety of other skeletal features were shared by many dinosaurs. However, because they were 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
Diapsid
Diapsids are a group of reptiles that developed two holes in each side of their skulls, about 300 million years ago during the late Carboniferous period. Living diapsids are extremely diverse, and include all crocodiles, lizards, snakes, and tuatara...

 reptiles, dinosaurs ancestrally had two pairs of temporal fenestrae (openings in the skull behind the eyes), and as members of the diapsid group Archosauria, had additional openings in the snout
Antorbital fenestra
An antorbital fenestra is an opening in the skull, in front of the eye sockets. This skull formation first appeared in archosaurs during the Triassic Period. Living birds today possess antorbital fenestrae, but the feature has been lost in modern crocodilians...

 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
Scapula
In anatomy, the scapula , omo, or shoulder blade, is the bone that connects the humerus with the clavicle ....

, or shoulder blade; a sacrum
Sacrum
In vertebrate anatomy the sacrum is a large, triangular bone at the base of the spine and at the upper and back part of the pelvic cavity, where it is inserted like a wedge between the two hip bones. Its upper part connects with the last lumbar vertebra, and bottom part with the coccyx...

 composed of three or more fused vertebrae (three are found in some other archosaurs, but only two are found in Herrerasaurus
Herrerasaurus
Herrerasaurus was one of the earliest dinosaurs. All known specimens of this carnivore have been discovered in rocks of late Ladinian age in northwestern Argentina...

); and an acetabulum
Acetabulum
The acetabulum is a concave surface of the pelvis. The head of the femur meets with the pelvis at the acetabulum, forming the hip joint.-Structure:...

, or hip socket, with a hole at the center of its inside surface (closed in Saturnalia
Saturnalia (dinosaur)
Saturnalia is an extinct genus of basal sauropodomorph dinosaur known from the Triassic of Rio Grande do Sul, southern Brazil.-Discovery:Saturnalia was originally named on the basis of three partial skeletons...

, for example). Another difficulty of determining distinctly dinosaurian features is that early dinosaurs and other archosaurs from the Late Triassic are often poorly known and were similar in many ways; these animals have sometimes been misidentified in the literature.

Dinosaurs stood erect in a manner similar to most modern mammals, but distinct from most other reptiles, whose limbs sprawl out to either side. Their posture was 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 dinosaurs to breathe easily while moving, which likely permitted stamina and activity levels that surpassed those of "sprawling" reptiles
Carrier's constraint
Carrier's constraint is the observation that air-breathing vertebrates which have two lungs and flex their bodies sideways during locomotion find it very difficult to move and breathe at the same time, because:...

. Erect limbs probably also helped support the evolution of large size by reducing bending stresses on limbs. Some non-dinosaurian archosaurs, including rauisuchia
Rauisuchia
Rauisuchia is a group of predatory and mostly large Triassic archosaurs. As a clade, Rauisuchia includes these Triassic forms and all crocodylomorphs, which are descendants of Triassic rauisuchians. The group in its traditional sense is paraphyletic, because it does not include crocodylomorph...

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
Ilium (bone)
The ilium is the uppermost and largest bone of the pelvis, and appears in most vertebrates including mammals and birds, but not bony fish. All reptiles have an ilium except snakes, although some snake species have a tiny bone which is considered to be an ilium.The name comes from the Latin ,...

 was rotated to form an overhanging shelf.

Origins and early evolution



Dinosaurs diverged from their archosaur
Archosaur
Archosaurs are a group of diapsid amniotes whose living representatives consist of modern birds and crocodilians. This group also includes all extinct non-avian dinosaurs, many extinct crocodilian relatives, and pterosaurs. Archosauria, the archosaur clade, is a crown group that includes the most...

 ancestors approximately 230 million years ago during the Middle to Late Triassic
Triassic
The Triassic is a geologic period and system that extends from about 250 to 200 Mya . As the first period of the Mesozoic Era, the Triassic follows the Permian and is followed by the Jurassic. Both the start and end of the Triassic are marked by major extinction events...

 period, roughly 20 million years after the Permian–Triassic extinction event wiped out an estimated 95% of all life on Earth
Life on Earth
Life on Earth: A Natural History by David Attenborough is a television natural history series made by the BBC in association with Warner Bros. and Reiner Moritz Productions...

. Radiometric dating
Radiometric dating
Radiometric dating is a technique used to date materials such as rocks, usually based on a comparison between the observed abundance of a naturally occurring radioactive isotope and its decay products, using known decay rates...

 of the rock formation
Rock formation
This is a list of rock formations that include isolated, scenic, or spectacular surface rock outcrops. These formations are usually the result of weathering and erosion sculpting the existing rock...

 that contained fossils from the early dinosaur genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...

 Eoraptor
Eoraptor
Eoraptor was one of the world's earliest dinosaurs. It was a two-legged saurischian, close to the ancestry of theropods and sauropodomorphs. It lived ca. 231.4 million years ago, in what is now the northwestern region of Argentina...

establishes its presence in the fossil record at this time. Paleontologists believe Eoraptor resembles the common ancestor
Common descent
In evolutionary biology, a group of organisms share common descent if they have a common ancestor. There is strong quantitative support for the theory that all living organisms on Earth are descended from a common ancestor....

 of all dinosaurs; if this is true, its traits suggest that the first dinosaurs were small, bipedal predators
Predation
In ecology, predation describes a biological interaction where a predator feeds on its prey . Predators may or may not kill their prey prior to feeding on them, but the act of predation always results in the death of its prey and the eventual absorption of the prey's tissue through consumption...

. The discovery of primitive, dinosaur-like ornithodirans such as Marasuchus
Marasuchus
Marasuchus is a genus of dinosaur-like ornithodiran from the middle Triassic Chañares Formation of Argentina. The species Marasuchus lilloensis was originally described as a second species of Lagosuchus, L. lilloensis...

and Lagerpeton
Lagerpeton
Lagerpeton is a genus of basal dinosauromorph from the Ladinian . Lagerpeton is known from several specimens of hindlimbs, hips, vertebrae, and feet. It was about 0.7 metres long and was found in the Chañares Formation of Argentina. It has unique feet, with an unusually long fourth toe...

in Argentinian
Argentina
Argentina , officially the Argentine Republic , is the second largest country in South America by land area, after Brazil. It is constituted as a federation of 23 provinces and an autonomous city, Buenos Aires...

 Middle Triassic
Middle Triassic
In the geologic timescale, the Middle Triassic is the second of three epochs of the Triassic period or the middle of three series in which the Triassic system is divided. It spans the time between 245 ± 1.5 Ma and 228 ± 2 Ma...

 strata supports this view; analysis of recovered fossils suggests that these animals were indeed small, bipedal predators.

When dinosaurs appeared, terrestrial habitats were occupied by various types of basal archosaurs and therapsids
Therapsida
Therapsida is a group of the most advanced synapsids, and include the ancestors of mammals. Many of the traits today seen as unique to mammals had their origin within early therapsids, including hair, lactation, and an erect posture. The earliest fossil attributed to Therapsida is believed to be...

, such as aetosaur
Aetosaur
Aetosaurs are an extinct order of heavily armoured, medium- to large-sized Late Triassic herbivorous archosaurs. They have small heads, upturned snouts, erect limbs, and a body covered by plate-like scutes. All aetosaurs belong to the family Stagonolepididae...

s, cynodont
Cynodont
Cynodontia or cynodonts are a taxon of therapsids which first appeared in the Late Permian and were eventually distributed throughout all seven continents by the Early Triassic . This clade includes modern mammals and their extinct close relatives. They were one of the most diverse groups of...

s, dicynodont
Dicynodont
Dicynodontia is a taxon of anomodont therapsids or mammal-like reptiles. Dicynodonts were small to large herbivorous animals with two tusks, hence their name, which means 'two dog tooth'...

s, ornithosuchids
Ornithosuchidae
Ornithosuchidae is an extinct family of reptiles from the Triassic period that were distantly related to crocodilians. They are classified as crurotarsan archosaurs. Ornithosuchids were quadrupedal and facultatively bipedal, meaning they had the ability to walk on two legs for short periods of time...

, rauisuchia
Rauisuchia
Rauisuchia is a group of predatory and mostly large Triassic archosaurs. As a clade, Rauisuchia includes these Triassic forms and all crocodylomorphs, which are descendants of Triassic rauisuchians. The group in its traditional sense is paraphyletic, because it does not include crocodylomorph...

s, and rhynchosaur
Rhynchosaur
Rhynchosaurs were a group of Triassic diapsid reptiles related to the archosaurs.-Description:Rhynchosaurs were herbivores, and at times abundant , with stocky bodies and a powerful beak...

s. Most of these other animals became extinct in the Triassic, in one of two events. First, at about the boundary between the Carnian
Carnian
The Carnian is the lowermost stage of the Upper Triassic series . It lasted from about 228.7 till 216.5 million years ago . The Carnian is preceded by the Ladinian and is followed by the Norian...

 and Norian
Norian
The Norian is a division of the Triassic geological period. It has the rank of an age or stage . The Norian lasted from 216.5 ± 2.0 to 203.6 ± 1.5 million years ago. It was preceded by the Carnian and succeeded by the Rhaetian.-Stratigraphic definitions:The Norian was named after the Noric Alps in...

 faunal stage
Faunal stage
In chronostratigraphy, a stage is a succession of rock strata laid down in a single age on the geologic timescale, which usually represents millions of years of deposition. A given stage of rock and the corresponding age of time will by convention have the same name, and the same boundaries.Rock...

s (about 215 million years ago), dicynodonts and a variety of basal archosauromorphs
Archosauromorpha
Archosauromorpha is an infraclass of diapsid reptiles that first appeared during the late Permian and became more common during the Triassic. Included in this infraclass are the groups Rhynchosauria, Trilophosauridae, Prolacertiformes, Archosauriformes, and, tentatively, Choristodera...

, including the prolacertiforms
Prolacertiformes
Prolacertiformes were an order of archosauromorph reptiles that lived during the Permian and Triassic Periods...

 and rhynchosaurs, became extinct. This was followed by the Triassic–Jurassic extinction event (about 200 million years ago), that saw the end of most of the other groups of early archosaurs, like aetosaurs, ornithosuchids, phytosaur
Phytosaur
Phytosaurs are an extinct group of large semi-aquatic Late Triassic archosaurs. Phytosaurs belong to the family Phytosauridae and the order Phytosauria. They were long-snouted and heavily armoured, bearing a remarkable resemblance to modern crocodiles in size, appearance, and lifestyle, an example...

s, and rauisuchians. These losses left behind a land fauna of crocodylomorphs
Crocodylomorpha
The Crocodylomorpha are an important group of archosaurs that include the crocodilians and their extinct relatives.During Mesozoic and early Tertiary times the Crocodylomorpha were far more diverse than they are now. Triassic forms were small, lightly built, active terrestrial animals. These were...

, dinosaurs, mammal
Mammal
Mammals are members of a class of air-breathing vertebrate animals characterised by the possession of endothermy, hair, three middle ear bones, and mammary glands functional in mothers with young...

s, pterosaurians, and turtle
Turtle
Turtles are reptiles of the order Testudines , characterised by a special bony or cartilaginous shell developed from their ribs that acts as a shield...

s. The first few lines of early dinosaurs diversified
Adaptive radiation
In evolutionary biology, adaptive radiation is the evolution of ecological and phenotypic diversity within a rapidly multiplying lineage. Starting with a recent single ancestor, this process results in the speciation and phenotypic adaptation of an array of species exhibiting different...

 through the Carnian
Carnian
The Carnian is the lowermost stage of the Upper Triassic series . It lasted from about 228.7 till 216.5 million years ago . The Carnian is preceded by the Ladinian and is followed by the Norian...

 and Norian
Norian
The Norian is a division of the Triassic geological period. It has the rank of an age or stage . The Norian lasted from 216.5 ± 2.0 to 203.6 ± 1.5 million years ago. It was preceded by the Carnian and succeeded by the Rhaetian.-Stratigraphic definitions:The Norian was named after the Noric Alps in...

 stages
Faunal stage
In chronostratigraphy, a stage is a succession of rock strata laid down in a single age on the geologic timescale, which usually represents millions of years of deposition. A given stage of rock and the corresponding age of time will by convention have the same name, and the same boundaries.Rock...

 of the Triassic, most likely by occupying the niches of the groups that became extinct.

Evolution and paleobiogeography



Dinosaur evolution after the Triassic follows changes in vegetation and the location of continents. In the Late Triassic and Early Jurassic, the continents were connected as the single landmass Pangaea
Pangaea
Pangaea, Pangæa, or Pangea is hypothesized as a supercontinent that existed during the Paleozoic and Mesozoic eras about 250 million years ago, before the component continents were separated into their current configuration....

, and there was a worldwide dinosaur fauna mostly composed of coelophysoid
Coelophysoidea
Coelophysoids were common dinosaurs of the Late Triassic and Early Jurassic periods. They were widespread geographically, probably living on all continents. Coelophysoids were all slender, carnivorous forms with a superficial similarity to the coelurosaurs, with which they were formerly...

 carnivore
Carnivore
A carnivore meaning 'meat eater' is an organism that derives its energy and nutrient requirements from a diet consisting mainly or exclusively of animal tissue, whether through predation or scavenging...

s and prosauropod herbivore
Herbivore
Herbivores are organisms that are anatomically and physiologically adapted to eat plant-based foods. Herbivory is a form of consumption in which an organism principally eats autotrophs such as plants, algae and photosynthesizing bacteria. More generally, organisms that feed on autotrophs in...

s. Gymnosperm
Gymnosperm
The gymnosperms are a group of seed-bearing plants that includes conifers, cycads, Ginkgo, and Gnetales. The term "gymnosperm" comes from the Greek word gymnospermos , meaning "naked seeds", after the unenclosed condition of their seeds...

 plants (particularly conifers), a potential food source, radiated in the Late Triassic. Prosauropods 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 ceratosauria
Ceratosauria
Ceratosaurs are members of a group of theropod dinosaurs defined as all theropods sharing a more recent common ancestry with Ceratosaurus than with birds. There is presently no universally agreed upon listing of species or diagnostic characters of Ceratosauria, though they were less derived...

ns, spinosauroids, and carnosauria
Carnosauria
Carnosauria is a group of large predatory dinosaurs that lived during the Jurassic and Cretaceous periods. While it originally contained a wide assortment of giant theropods that were not closely related, the group has since been defined to encompass only the allosaurs and their closest kin...

ns, and herbivores consisting of stegosauria
Stegosauria
Known colloquially as stegosaurs, the Stegosauria are a group of herbivorous dinosaurs of the Jurassic and Early Cretaceous Periods, being found mostly in the Northern Hemisphere, predominantly in what is now North America and China....

n ornithischians and large sauropods. Examples of this include the Morrison Formation
Morrison Formation
The Morrison Formation is a distinctive sequence of Late Jurassic sedimentary rock that is found in the western United States, which has been the most fertile source of dinosaur fossils in North America. It is composed of mudstone, sandstone, siltstone and limestone and is light grey, greenish...

 of North America and Tendaguru Beds
Tendaguru
The Tendaguru Beds are a fossil-rich formation in Tanzania. It has been considered the richest of Late Jurassic strata in Africa. Continental reconstructions show Tendaguru to have been in the southern hemisphere during the Late Jurassic. Tendaguru is similar to the Morrison Formation except in...

 of Tanzania. Dinosaurs in China show some differences, with specialized sinraptorid
Sinraptoridae
Sinraptorids were a family of carnivorous theropod dinosaurs. They tended to be large predators, some growing to sizes of . Sinraptorids are carnosaurs, and many were initially classified within Megalosauridae or Allosauridae prior to recent analysis...

 theropods and unusual, long-necked sauropods like Mamenchisaurus
Mamenchisaurus
Mamenchisaurus was a plant-eating four-legged dinosaur, known for its remarkably long neck which made up half its total length. It is known from numerous species which ranged in time from 160 to 145 million years ago, from the Oxfordian to Tithonian ages of the late Jurassic Period of...

. Ankylosauria
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. Ankylosaurs were bulky quadrupeds, with short, powerful limbs. They are first known to have appeared in the early Jurassic Period of...

ns and ornithopods were also becoming more common, but prosauropods had become extinct. Conifers and pteridophyte
Pteridophyte
The pteridophytes are vascular plants that produce neither flowers nor seeds, and are hence called vascular cryptogams. Instead, they reproduce and disperse only via spores. Pteridophytes include horsetails, ferns, club mosses, and quillworts...

s were the most common plants. Sauropods, like the earlier prosauropods, were not oral processors, but ornithischians were evolving various means of dealing with food in the mouth, including potential cheek
Cheek
Cheeks constitute the area of the face below the eyes and between the nose and the left or right ear. They may also be referred to as jowls. "Buccal" means relating to the cheek. In humans, the region is innervated by the buccal nerve...

-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 maniraptora
Maniraptora
Maniraptora is a clade of coelurosaurian dinosaurs which includes the birds and the dinosaurs that were more closely related to them than to Ornithomimus velox. It contains the major subgroups Avialae, Deinonychosauria, Oviraptorosauria and Therizinosauria. Ornitholestes and the Alvarezsauroidea...

n coelurosauria
Coelurosauria
Coelurosauria is the clade containing all theropod dinosaurs more closely related to birds than to carnosaurs. In the past, it was used to refer to all small theropods, although this classification has been abolished...

ns.

By the Early Cretaceous and the ongoing breakup of Pangaea, dinosaurs were becoming strongly differentiated by landmass. The earliest part of this time saw the spread of ankylosaurians, iguanodontians, and brachiosaurids
Brachiosauridae
Brachiosauridae are a family of dinosaurs, whose members are known as brachiosaurids. They were herbivorous quadrupeds with longer forelegs than hind legs - the name derives from the Greek for arm lizard - and long necks...

 through Europe, North America, and northern Africa. These were later supplemented or replaced in Africa by large spinosaurid and carcharodontosaurid
Carcharodontosauridae
Carcharodontosaurids were a group of carnivorous theropod dinosaurs. In 1931 Ernst Stromer named Carcharodontosauridae as a family, in modern paleontology this name indicates a clade within Carnosauria...

 theropods, and rebbachisaurid
Rebbachisauridae
Rebbachisauridae is a family of sauropod dinosaurs known from fragmentary fossil remains from the Cretaceous of South America, Africa, and Europe.-Taxonomy:...

 and titanosaurian sauropods, also found in South America. In Asia, maniraptora
Maniraptora
Maniraptora is a clade of coelurosaurian dinosaurs which includes the birds and the dinosaurs that were more closely related to them than to Ornithomimus velox. It contains the major subgroups Avialae, Deinonychosauria, Oviraptorosauria and Therizinosauria. Ornitholestes and the Alvarezsauroidea...

n coelurosauria
Coelurosauria
Coelurosauria is the clade containing all theropod dinosaurs more closely related to birds than to carnosaurs. In the past, it was used to refer to all small theropods, although this classification has been abolished...

ns like dromaeosaurids
Dromaeosauridae
Dromaeosauridae is a family of bird-like theropod dinosaurs. They were small- to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from Greek dromeus meaning 'runner' and sauros meaning 'lizard'...

, troodontids
Troodontidae
Troodontidae is a family of bird-like theropod dinosaurs. During most of the 20th century, troodontid fossils were few and scrappy and they have therefore been allied, at various times, with many dinosaurian lineages...

, and oviraptorosauria
Oviraptorosauria
Oviraptorosaurs are a group of feathered maniraptoran dinosaurs from the Cretaceous Period of what are now Asia and North America. They are distinct for their characteristically short, beaked, parrot - like skulls, with or without bony crests atop the head...

ns became the common theropods, and ankylosaurids
Ankylosauridae
An ankylosaurid is a member of the Ankylosauridae family of armored dinosaurs that evolved 125 million years ago and became extinct 65 million years ago during the Cretaceous-Tertiary extinction event...

 and early ceratopsia
Ceratopsia
Ceratopsia or Ceratopia is a group of herbivorous, beaked dinosaurs which thrived in what are now North America, Europe, and Asia, during the Cretaceous Period, although ancestral forms lived earlier, in the Jurassic. The earliest known ceratopsian, Yinlong downsi, lived between 161.2 and 155.7...

ns like Psittacosaurus became important herbivores. Meanwhile, Australia was home to a fauna of basal ankylosaurians, hypsilophodont
Hypsilophodont
Hypsilophodonts were small ornithopod dinosaurs, regarded as fast, herbivorous bipeds on the order of 1–2 meters long . They are known from Asia, Australia, Europe, New Zealand, North America, and South America, from rocks of Middle Jurassic to late Cretaceous age...

s, 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 tooth batteries, taken to its extreme in hadrosaurid
Hadrosaurid
Hadrosaurids or duck-billed dinosaurs are members of the family Hadrosauridae, and include ornithopods such as Edmontosaurus and Parasaurolophus. They were common herbivores in the Upper Cretaceous Period of what are now Asia, Europe and North America. They are descendants of the Upper...

s. Some sauropods also evolved tooth batteries, best exemplified by the rebbachisaurid Nigersaurus
Nigersaurus
Nigersaurus is a genus of diplodocoid sauropod dinosaur from the middle Cretaceous period, about 119 to 99 million years ago during the Aptian or Albian age. This dinosaur was described by Paul Sereno and colleagues in 1999...

.

There were three general dinosaur faunas in the Late Cretaceous. In the northern continents of North America and Asia, the major theropods were tyrannosaurids
Tyrannosauridae
Tyrannosauridae is a family of coelurosaurian theropod dinosaurs which comprises two subfamilies containing up to six genera, including the eponymous Tyrannosaurus. The exact number of genera is controversial, with some experts recognizing as few as three...

 and various types of smaller maniraptoran theropods, with a predominantly ornithischian herbivore assemblage of hadrosaurids, ceratopsians, ankylosaurids, and pachycephalosauria
Pachycephalosauria
Pachycephalosauria is a clade of ornithischian dinosaurs. Well-known genera include Pachycephalosaurus, Stegoceras, Stygimoloch, and Dracorex. Most lived during the Late Cretaceous Period, in what is now North America and Asia. They were all bipedal, herbivorous/omnivorous animals with thick skulls...

ns. In the southern continents that had made up the now-splitting Gondwana
Gondwana
In paleogeography, Gondwana , originally Gondwanaland, was the southernmost of two supercontinents that later became parts of the Pangaea supercontinent. It existed from approximately 510 to 180 million years ago . Gondwana is believed to have sutured between ca. 570 and 510 Mya,...

, abelisaurids
Abelisauridae
Abelisauridae is a family of ceratosaurian theropod dinosaurs. Abelisaurids thrived during the Cretaceous Period, on the ancient southern supercontinent of Gondwana, and today their fossil remains are found on the modern continents of Africa and South America, as well as on the Indian...

 were the common theropods, and titanosaurian sauropods the common herbivores. Finally, in Europe, dromaeosaurids, rhabdodontid
Rhabdodontidae
Rhabdodontids were herbivorous ornithopod dinosaurs from the Cretaceous Period. Rhabdodontids were similar to large, robust hypsilophodonts, with deep skulls and jaws. The family was first proposed by David B. Weishampel et al. in 2002...

 iguanodontians, nodosaurid
Nodosauridae
Nodosauridae is a family of ankylosaurian dinosaurs, from the Late Jurassic to the Late Cretaceous Period of what are now North America, Asia, Antarctica and Europe.-Characteristics:...

 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 extremely diverse across North America and Asia. Theropods were also radiating as herbivores or omnivores, with therizinosaur
Therizinosaur
Therizinosaurs are theropod dinosaurs belonging to the clade Therizinosauria. Therizinosaur fossils have been found in Early through Late Cretaceous deposits in Mongolia, the People's Republic of China and Western North America...

ians and ornithomimosauria
Ornithomimosauria
The Ornithomimosauria, ornithomimosaurs or ostrich dinosaurs were theropod dinosaurs which bore a superficial resemblance to modern ostriches. They were fast, omnivorous or herbivorous dinosaurs from the Cretaceous Period of Laurasia...

ns becoming common.

The Cretaceous–Paleogene extinction event, which occurred approximately 65 million years ago at the end of the Cretaceous period, caused the extinction of all dinosaurs except for the birds. Some other diapsid
Diapsid
Diapsids are a group of reptiles that developed two holes in each side of their skulls, about 300 million years ago during the late Carboniferous period. Living diapsids are extremely diverse, and include all crocodiles, lizards, snakes, and tuatara...

 groups, such as crocodilians, lizard
Lizard
Lizards are a widespread group of squamate reptiles, with nearly 3800 species, ranging across all continents except Antarctica as well as most oceanic island chains...

s, snake
Snake
Snakes are elongate, legless, carnivorous reptiles of the suborder Serpentes that can be distinguished from legless lizards by their lack of eyelids and external ears. Like all squamates, snakes are ectothermic, amniote vertebrates covered in overlapping scales...

s, sphenodontia
Sphenodontia
Sphenodontia is an order of lizard-like reptiles that includes only one living genus, the tuatara , and only two living species...

ns, and choristodera
Choristodera
Choristodera is an order of semi-aquatic diapsid reptiles which ranged from the Middle Jurassic, or possibly Late Triassic, to at least the early Miocene. Choristoderes have been found in North America, Asia, and Europe. The most common fossils are typically found from the Late Cretaceous to the...

ns, also survived the event.

Classification


Dinosaurs (including birds) are archosaur
Archosaur
Archosaurs are a group of diapsid amniotes whose living representatives consist of modern birds and crocodilians. This group also includes all extinct non-avian dinosaurs, many extinct crocodilian relatives, and pterosaurs. Archosauria, the archosaur clade, is a crown group that includes the most...

s, like modern crocodilia
Crocodilia
Crocodilia is an order of large reptiles that appeared about 84 million years ago in the late Cretaceous Period . They are the closest living relatives of birds, as the two groups are the only known survivors of the Archosauria...

ns. 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 are usually regarded as a superorder or an unranked clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...

. They are divided into two orders
Order (biology)
In scientific classification used in biology, the order is# a taxonomic rank used in the classification of organisms. Other well-known ranks are life, domain, kingdom, phylum, class, family, genus, and species, with order fitting in between class and family...

, Saurischia
Saurischia
Saurischia meaning 'lizard' and ischion meaning 'hip joint') is one of the two orders, or basic divisions, of dinosaurs. In 1888, Harry Seeley classified dinosaurs into two orders, based on their hip structure...

 and Ornithischia
Ornithischia
Ornithischia or Predentata is an extinct order of beaked, herbivorous dinosaurs. The name ornithischia is derived from the Greek ornitheos meaning 'of a bird' and ischion meaning 'hip joint'...

, depending upon pelvic
Pelvis
In human anatomy, the pelvis is the lower part of the trunk, between the abdomen and the lower limbs .The pelvis includes several structures:...

 structure. Saurischia includes those taxa sharing a more recent common ancestor with birds than with Ornithischia, while Ornithischia includes all taxa
Taxon
|thumb|270px|[[African elephants]] form a widely-accepted taxon, the [[genus]] LoxodontaA taxon is a group of organisms, which a taxonomist adjudges to be a unit. Usually a taxon is given a name and a rank, although neither is a requirement...

 sharing a more recent common ancestor with Triceratops
Triceratops
Triceratops is a genus of herbivorous ceratopsid dinosaur which lived during the late Maastrichtian stage of the Late Cretaceous Period, around 68 to 65 million years ago in what is now North America. It was one of the last dinosaur genera to appear before the great Cretaceous–Paleogene...

than with Saurischia. Saurischians ("lizard-hipped", from the Greek
Ancient Greek
Ancient Greek is the stage of the Greek language in the periods spanning the times c. 9th–6th centuries BC, , c. 5th–4th centuries BC , and the c. 3rd century BC – 6th century AD of ancient Greece and the ancient world; being predated in the 2nd millennium BC by Mycenaean Greek...

 sauros (σαυρος) meaning "lizard" and ischion (ισχιον) meaning "hip joint") retained the hip structure of their ancestors, with a pubis
Pubis (bone)
In vertebrates, the pubic bone is the ventral and anterior of the three principal bones composing either half of the pelvis.It is covered by a layer of fat, which is covered by the mons pubis....

 bone directed cranially
Anatomical terms of location
Standard anatomical terms of location are designations employed in science that deal with the anatomy of animals to avoid ambiguities that might otherwise arise. They are not language-specific, and thus require no translation...

, or forward. This basic form was modified by rotating the pubis backward to varying degrees in several groups (Herrerasaurus
Herrerasaurus
Herrerasaurus was one of the earliest dinosaurs. All known specimens of this carnivore have been discovered in rocks of late Ladinian age in northwestern Argentina...

, therizinosaur
Therizinosaur
Therizinosaurs are theropod dinosaurs belonging to the clade Therizinosauria. Therizinosaur fossils have been found in Early through Late Cretaceous deposits in Mongolia, the People's Republic of China and Western North America...

oids, dromaeosaurids
Dromaeosauridae
Dromaeosauridae is a family of bird-like theropod dinosaurs. They were small- to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from Greek dromeus meaning 'runner' and sauros meaning 'lizard'...

, and bird
Bird
Birds are feathered, winged, bipedal, endothermic , egg-laying, vertebrate animals. Around 10,000 living species and 188 families makes them the most speciose class of tetrapod vertebrates. They inhabit ecosystems across the globe, from the Arctic to the Antarctic. Extant birds range in size from...

s). Saurischia includes the theropods
Theropoda
Theropoda is both a suborder of bipedal saurischian dinosaurs, and a clade consisting of that suborder and its descendants . Dinosaurs belonging to the suborder theropoda were primarily carnivorous, although a number of theropod groups evolved herbivory, omnivory, and insectivory...

 (bipedal and mostly carnivores, except for birds) and sauropodomorphs
Sauropodomorpha
Sauropodomorpha is an extinct clade of long-necked, herbivorous, saurischian dinosaurs which includes the sauropods and their ancestral relatives. Sauropods generally grew to very large sizes, had long necks and tails, were quadrupedal, and became the largest animals to ever walk the Earth. The...

 (long-necked quadrupedal herbivore
Herbivore
Herbivores are organisms that are anatomically and physiologically adapted to eat plant-based foods. Herbivory is a form of consumption in which an organism principally eats autotrophs such as plants, algae and photosynthesizing bacteria. More generally, organisms that feed on autotrophs in...

s).

By contrast, ornithischians ("bird-hipped", from the Greek
Ancient Greek
Ancient Greek is the stage of the Greek language in the periods spanning the times c. 9th–6th centuries BC, , c. 5th–4th centuries BC , and the c. 3rd century BC – 6th century AD of ancient Greece and the ancient world; being predated in the 2nd millennium BC by Mycenaean Greek...

 ornitheios (ορνιθειος) meaning "of a bird" and ischion (ισχιον) meaning "hip joint") had a pelvis that superficially resembled a bird's pelvis: the pubis
Pubis (bone)
In vertebrates, the pubic bone is the ventral and anterior of the three principal bones composing either half of the pelvis.It is covered by a layer of fat, which is covered by the mons pubis....

 bone was oriented caudally
Anatomical terms of location
Standard anatomical terms of location are designations employed in science that deal with the anatomy of animals to avoid ambiguities that might otherwise arise. They are not language-specific, and thus require no translation...

 (rear-pointing) Unlike birds, the ornithischian pubis also usually had an additional forward-pointing process. Ornithischia includes a variety of herbivores. (NB: the terms "lizard hip" and "bird hip" are misnomers – birds evolved from dinosaurs with "lizard hips".)


Taxonomy


The following is a simplified classification of dinosaur groups based on their evolutionary relationships. A more detailed version can be found at Dinosaur classification
Dinosaur classification
Dinosaur classification began in 1842 when Sir Richard Owen placed Iguanodon, Megalosaurus, and Hylaeosaurus in "a distinct tribe or suborder of Saurian Reptiles, for which I would propose the name of Dinosauria." In 1887 and 1888 Harry Seeley divided dinosaurs into the two orders Saurischia and...

.
The cross (†) is used to signify groups with no living members.



  • Dinosauria
  • Saurischia
    Saurischia
    Saurischia meaning 'lizard' and ischion meaning 'hip joint') is one of the two orders, or basic divisions, of dinosaurs. In 1888, Harry Seeley classified dinosaurs into two orders, based on their hip structure...

    ("lizard-hipped"; includes Theropoda and Sauropodomorpha)
  • Herrerasauridae
    Herrerasauridae
    Herrerasaurids are among the oldest known dinosaurs, appearing in the fossil record 231.4 million years ago . They became extinct by the end of the Triassic period. Herrerasaurids were small-sized carnivorous theropods or basal saurischians...

     (early bipedal carnivores)
  • Theropoda
    Theropoda
    Theropoda is both a suborder of bipedal saurischian dinosaurs, and a clade consisting of that suborder and its descendants . Dinosaurs belonging to the suborder theropoda were primarily carnivorous, although a number of theropod groups evolved herbivory, omnivory, and insectivory...

    (all bipedal; most were carnivorous)
  • Coelophysoidea
    Coelophysoidea
    Coelophysoids were common dinosaurs of the Late Triassic and Early Jurassic periods. They were widespread geographically, probably living on all continents. Coelophysoids were all slender, carnivorous forms with a superficial similarity to the coelurosaurs, with which they were formerly...

     (small, early theropods; includes Coelophysis
    Coelophysis
    Coelophysis , meaning "hollow form" in reference to its hollow bones , is one of the earliest known genera of dinosaur...

    and close relatives)
  • Ceratosauria
    Ceratosauria
    Ceratosaurs are members of a group of theropod dinosaurs defined as all theropods sharing a more recent common ancestry with Ceratosaurus than with birds. There is presently no universally agreed upon listing of species or diagnostic characters of Ceratosauria, though they were less derived...

     (Ceratosaurus
    Ceratosaurus
    Ceratosaurus meaning "horned lizard", in reference to the horn on its nose , was a large predatory theropod dinosaur from the Late Jurassic Period , found in the Morrison Formation of North America, in Tanzania and Portugal...

    and close relatives, such as abelisaur
    Abelisaur
    Abelisaurs were a group of ceratosaurian dinosaurs which lived in the Southern hemisphere during the Cretaceous period. Some well-known dinosaurs of this group include Abelisaurus, Carnotaurus, and Majungasaurus. They are known for their small arms...

    ids)
  • Tetanurae
    Tetanurae
    Tetanurae, or "stiff tails", is a clade that includes most theropod dinosaurs, as well as birds. Tetanurans first appear during the early or middle Jurassic Period.-Definition:...

     ("stiff tails"; includes most theropods)
  • Megalosauroidea
    Megalosauroidea
    Megalosauroidea is a group of tetanuran theropod dinosaurs that lived from the Middle Jurassic to the Late Cretaceous period.-Classification:...

     (Megalosaurus
    Megalosaurus
    Megalosaurus is a genus of large meat-eating theropod dinosaurs of the Middle Jurassic period of Europe...

    , Spinosaurus
    Spinosaurus
    Spinosaurus is a genus of theropod dinosaur which lived in what is now North Africa, from the lower Albian to lower Cenomanian stages of the Cretaceous period, about 112 to 97 million years ago. This genus was first known from Egyptian remains discovered in 1912 and described by German...

    , and close relatives)
  • Carnosauria
    Carnosauria
    Carnosauria is a group of large predatory dinosaurs that lived during the Jurassic and Cretaceous periods. While it originally contained a wide assortment of giant theropods that were not closely related, the group has since been defined to encompass only the allosaurs and their closest kin...

     (Allosaurus
    Allosaurus
    Allosaurus is a genus of large theropod dinosaur that lived 155 to 150 million years ago during the late Jurassic period . The name Allosaurus means "different lizard". It is derived from the Greek /allos and /sauros...

    and close relatives, like Carcharodontosaurus
    Carcharodontosaurus
    Carcharodontosaurus was a gigantic carnivorous carcharodontosaurid dinosaur that lived around 100 to 93 million years ago, during the late Albian to early Cenomanian stages of the mid-Cretaceous Period...

    )
  • Coelurosauria
    Coelurosauria
    Coelurosauria is the clade containing all theropod dinosaurs more closely related to birds than to carnosaurs. In the past, it was used to refer to all small theropods, although this classification has been abolished...

     (closely related to birds; diverse, with a range of body sizes and niches)
  • Tyrannosauridae
    Tyrannosauridae
    Tyrannosauridae is a family of coelurosaurian theropod dinosaurs which comprises two subfamilies containing up to six genera, including the eponymous Tyrannosaurus. The exact number of genera is controversial, with some experts recognizing as few as three...

     (Tyrannosaurus
    Tyrannosaurus
    Tyrannosaurus meaning "tyrant," and sauros meaning "lizard") is a genus of coelurosaurian theropod dinosaur. The species Tyrannosaurus rex , commonly abbreviated to T. rex, is a fixture in popular culture. It lived throughout what is now western North America, with a much wider range than other...

    and close relatives; had reduced forelimbs)
  • Ornithomimosauria
    Ornithomimosauria
    The Ornithomimosauria, ornithomimosaurs or ostrich dinosaurs were theropod dinosaurs which bore a superficial resemblance to modern ostriches. They were fast, omnivorous or herbivorous dinosaurs from the Cretaceous Period of Laurasia...

     ("ostrich
    Ostrich
    The Ostrich is one or two species of large flightless birds native to Africa, the only living member of the genus Struthio. Some analyses indicate that the Somali Ostrich may be better considered a full species apart from the Common Ostrich, but most taxonomists consider it to be a...

    -mimics"; mostly toothless; carnivores to possible herbivores)
  • Maniraptora
    Maniraptora
    Maniraptora is a clade of coelurosaurian dinosaurs which includes the birds and the dinosaurs that were more closely related to them than to Ornithomimus velox. It contains the major subgroups Avialae, Deinonychosauria, Oviraptorosauria and Therizinosauria. Ornitholestes and the Alvarezsauroidea...

     ("hand snatchers"; had long, slender arms and fingers)
  • †Therizinosauria (bipedal herbivores with large hand claws and small heads)
  • Oviraptorosauria
    Oviraptorosauria
    Oviraptorosaurs are a group of feathered maniraptoran dinosaurs from the Cretaceous Period of what are now Asia and North America. They are distinct for their characteristically short, beaked, parrot - like skulls, with or without bony crests atop the head...

     (mostly toothless; their diet and lifestyle are uncertain)
  • Deinonychosauria
    Deinonychosauria
    The Deinonychosauria were a clade of coelurosaurian theropod dinosaurs from the Late Jurassic and Cretaceous periods. These omnivores and carnivores are known for their switchblade-like second toe claws and for displaying numerous bird-like characteristics. The clade has been divided into two...

     (small- to medium-sized; bird-like, with a distinctive toe claw)
  • Avialae
    Avialae
    Avialae is a clade of dinosaurs containing their only living representatives, birds , and the most immediate extinct relatives of birds.-Competing definitions:...

     (flying, feathered dinosaurs; includes Aves)
  • Sauropodomorpha
    Sauropodomorpha
    Sauropodomorpha is an extinct clade of long-necked, herbivorous, saurischian dinosaurs which includes the sauropods and their ancestral relatives. Sauropods generally grew to very large sizes, had long necks and tails, were quadrupedal, and became the largest animals to ever walk the Earth. The...

    (herbivores with small heads, long necks, long tails)
  • †Prosauropoda (early relatives of sauropods; small- to medium-sized; some possibly omnivorous; bipeds and quadrupeds)
  • Sauropoda
    Sauropoda
    Sauropoda , or the sauropods , are an infraorder of saurischian dinosaurs. They had long necks, long tails, small heads , and thick, pillar-like legs. They are notable for the enormous sizes attained by some species, and the group includes the largest animals to have ever lived on land...

     (very large and heavy, usually over 15 meters (49 feet) long; quadrupedal)
  • Diplodocoidea
    Diplodocoidea
    Diplodocoidea was a superfamily of sauropod dinosaurs, which included some of the longest animals of all time, including slender giants like Supersaurus, Diplodocus, Apatosaurus, and Amphicoelias...

     (skulls and tails elongated; teeth typically narrow and pencil-like)
  • Macronaria
    Macronaria
    Macronaria is a clade of sauropod dinosaurs from the Middle Jurassic to Late Cretaceous Periods of what are now North America, South America, Europe, Asia and Africa. The name means 'large nostrils' , in reference to the large nasal openings high on the head that probably supported fleshy...

     (boxy skulls; spoon- or pencil-shaped teeth)
  • Brachiosauridae
    Brachiosauridae
    Brachiosauridae are a family of dinosaurs, whose members are known as brachiosaurids. They were herbivorous quadrupeds with longer forelegs than hind legs - the name derives from the Greek for arm lizard - and long necks...

     (very long necks; forelimbs longer than hindlimbs)
  • †Titanosauria (diverse; stocky, with wide hips; most common in the Late Cretaceous of southern continents)

  • Ornithischia
    Ornithischia
    Ornithischia or Predentata is an extinct order of beaked, herbivorous dinosaurs. The name ornithischia is derived from the Greek ornitheos meaning 'of a bird' and ischion meaning 'hip joint'...

    ("bird-hipped"; diverse bipedal and quadrupedal herbivores)
  • Heterodontosauridae
    Heterodontosauridae
    Heterodontosauridae is a family of early ornithischian dinosaurs that were likely among the most basal members of the group...

     (small basal ornithopod herbivores/omnivores with prominent canine teeth)
  • Thyreophora
    Thyreophora
    The Thyreophora were a subgroup of the ornithischian dinosaurs...

    (armored dinosaurs; mostly quadrupeds)
  • Ankylosauria
    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. Ankylosaurs were bulky quadrupeds, with short, powerful limbs. They are first known to have appeared in the early Jurassic Period of...

     (scute
    Scute
    A scute or scutum is a bony external plate or scale, as on the shell of a turtle, the skin of crocodilians, the feet of some birds or the anterior portion of the mesonotum in insects.-Properties:...

    s as primary armor; some had club-like tails)
  • Stegosauria
    Stegosauria
    Known colloquially as stegosaurs, the Stegosauria are a group of herbivorous dinosaurs of the Jurassic and Early Cretaceous Periods, being found mostly in the Northern Hemisphere, predominantly in what is now North America and China....

     (spikes and plates as primary armor)
  • Ornithopoda (various sizes; bipeds and quadrupeds; evolved a method of chewing using skull flexibility and numerous teeth)
  • †Iguanodontia (herbivores which developed complex chewing mechanisms)
  • †Hadrosauridae (duck-billed dinosaurs)
  • Marginocephalia
    Marginocephalia
    Marginocephalia is a clade of ornithischian dinosaurs that includes the thick-skulled pachycephalosaurids, and horned ceratopsians. They were all herbivores, walking on two or four legs, and are characterized by a bony ridge or frill the back of the skull...

    (characterized by a cranial growth)
  • Pachycephalosauria
    Pachycephalosauria
    Pachycephalosauria is a clade of ornithischian dinosaurs. Well-known genera include Pachycephalosaurus, Stegoceras, Stygimoloch, and Dracorex. Most lived during the Late Cretaceous Period, in what is now North America and Asia. They were all bipedal, herbivorous/omnivorous animals with thick skulls...

     (bipeds with domed or knobby growth on skulls)
  • Ceratopsia
    Ceratopsia
    Ceratopsia or Ceratopia is a group of herbivorous, beaked dinosaurs which thrived in what are now North America, Europe, and Asia, during the Cretaceous Period, although ancestral forms lived earlier, in the Jurassic. The earliest known ceratopsian, Yinlong downsi, lived between 161.2 and 155.7...

     (quadrupeds with frills; many also had horns)

Paleobiology


Knowledge about dinosaurs is derived from a variety of fossil and non-fossil records, including fossilized bone
Bone
Bones are rigid organs that constitute part of the endoskeleton of vertebrates. They support, and protect the various organs of the body, produce red and white blood cells and store minerals. Bone tissue is a type of dense connective tissue...

s, feces
Feces
Feces, faeces, or fæces is a waste product from an animal's digestive tract expelled through the anus or cloaca during defecation.-Etymology:...

, trackway
Trackway
A trackway is an ancient route of travel for people or animals. In biology, a trackway can be a set of impressions in the soft earth, usually a set of footprints, left by an animal. A fossil trackway is the fossilized imprint of a trackway. Trackways have been found all over the world...

s, gastrolith
Gastrolith
A gastrolith, also called a stomach stone or gizzard stones, is a rock held inside a gastrointestinal tract. Gastroliths are retained in the muscular gizzard and used to grind food in animals lacking suitable grinding teeth. The grain size depends upon the size of the animal and the gastrolith's...

s, feather
Feather
Feathers are one of the epidermal growths that form the distinctive outer covering, or plumage, on birds and some non-avian theropod dinosaurs. They are considered the most complex integumentary structures found in vertebrates, and indeed a premier example of a complex evolutionary novelty. They...

s, impressions of skin, internal organs
Viscus
In anatomy, a viscus is an internal organ, and viscera is the plural form. The viscera, when removed from a butchered animal, are known collectively as offal...

 and soft tissue
Soft tissue
In anatomy, the term soft tissue refers to tissues that connect, support, or surround other structures and organs of the body, not being bone. Soft tissue includes tendons, ligaments, fascia, skin, fibrous tissues, fat, and synovial membranes , and muscles, nerves and blood vessels .It is sometimes...

s. Many fields of study contribute to our understanding of dinosaurs, including physics
Physics
Physics is a natural science that involves the study of matter and its motion through spacetime, along with related concepts such as energy and force. More broadly, it is the general analysis of nature, conducted in order to understand how the universe behaves.Physics is one of the oldest academic...

 (especially biomechanics
Biomechanics
Biomechanics is the application of mechanical principles to biological systems, such as humans, animals, plants, organs, and cells. Perhaps one of the best definitions was provided by Herbert Hatze in 1974: "Biomechanics is the study of the structure and function of biological systems by means of...

), chemistry
Chemistry
Chemistry is the science of matter, especially its chemical reactions, but also its composition, structure and properties. Chemistry is concerned with atoms and their interactions with other atoms, and particularly with the properties of chemical bonds....

, biology
Biology
Biology is a natural science concerned with the study of life and living organisms, including their structure, function, growth, origin, evolution, distribution, and taxonomy. Biology is a vast subject containing many subdivisions, topics, and disciplines...

, and the earth sciences (of which paleontology
Paleontology
Paleontology "old, ancient", ὄν, ὀντ- "being, creature", and λόγος "speech, thought") is the study of prehistoric life. It includes the study of fossils to determine organisms' evolution and interactions with each other and their environments...

 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 periods. Theropod dinosaurs, when sorted by estimated weight into categories based on order of magnitude
Order of magnitude
An order of magnitude is the class of scale or magnitude of any amount, where each class contains values of a fixed ratio to the class preceding it. In its most common usage, the amount being scaled is 10 and the scale is the exponent being applied to this amount...

, most often fall into the 100 to 1000 kilogram (220 to 2200 lb) category, whereas recent
Holocene
The Holocene is a geological epoch which began at the end of the Pleistocene and continues to the present. The Holocene is part of the Quaternary period. Its name comes from the Greek words and , meaning "entirely recent"...

 predatory carnivora
Carnivora
The diverse order Carnivora |Latin]] carō "flesh", + vorāre "to devour") includes over 260 species of placental mammals. Its members are formally referred to as carnivorans, while the word "carnivore" can refer to any meat-eating animal...

ns peak in the 10 to 100 kilogram (22 to 220 lb) category. The mode
Mode (statistics)
In statistics, the mode is the value that occurs most frequently in a data set or a probability distribution. In some fields, notably education, sample data are often called scores, and the sample mode is known as the modal score....

 of dinosaur body masses is between one and ten metric tonnes. This contrasts sharply with the size of Cenozoic
Cenozoic
The Cenozoic era is the current and most recent of the three Phanerozoic geological eras and covers the period from 65.5 mya to the present. The era began in the wake of the Cretaceous–Tertiary extinction event at the end of the Cretaceous that saw the demise of the last non-avian dinosaurs and...

 mammals, estimated by the National Museum of Natural History
National Museum of Natural History
The National Museum of Natural History is a natural history museum administered by the Smithsonian Institution, located on the National Mall in Washington, D.C., United States. Admission is free and the museum is open 364 days a year....

 as about 2 to 5 kilograms (5 to 10 lb).

The sauropods
Sauropoda
Sauropoda , or the sauropods , are an infraorder of saurischian dinosaurs. They had long necks, long tails, small heads , and thick, pillar-like legs. They are notable for the enormous sizes attained by some species, and the group includes the largest animals to have ever lived on land...

 were the largest and heaviest dinosaurs. For much of the dinosaur era, the smallest sauropods were larger than anything else in their habitat, and the largest were an order of magnitude
Order of magnitude
An order of magnitude is the class of scale or magnitude of any amount, where each class contains values of a fixed ratio to the class preceding it. In its most common usage, the amount being scaled is 10 and the scale is the exponent being applied to this amount...

 more massive than anything else that has since walked the Earth. Giant prehistoric mammal
Mammal
Mammals are members of a class of air-breathing vertebrate animals characterised by the possession of endothermy, hair, three middle ear bones, and mammary glands functional in mothers with young...

s such as the Paraceratherium
Paraceratherium
Paraceratherium, also commonly known as Indricotherium or Baluchitherium , is an extinct genus of gigantic hornless rhinoceros-like mammals of the family Hyracodontidae, endemic to Eurasia and Asia during the Eocene to Oligocene 37.2—23.030 Mya, existing for approximately...

and the Columbian mammoth
Mammoth
A mammoth is any species of the extinct genus Mammuthus. These proboscideans are members of Elephantidae, the family of elephants and mammoths, and close relatives of modern elephants. They were often equipped with long curved tusks and, in northern species, a covering of long hair...

 were dwarfed by the giant sauropods, and only a handful of modern aquatic animals approach or surpass them in size – most notably the blue whale
Blue Whale
The blue whale is a marine mammal belonging to the suborder of baleen whales . At in length and or more in weight, it is the largest known animal to have ever existed....

, which reaches up to kg ( lb) and over 30 metres (98.4 ft) in length. 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 formation
Rock formation
This is a list of rock formations that include isolated, scenic, or spectacular surface rock outcrops. These formations are usually the result of weathering and erosion sculpting the existing rock...

s 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 and smallest dinosaurs. This is because only a tiny percentage of animals ever fossilize, and most of these remain buried in the earth. Few of the specimens that are recovered are complete skeletons, and impressions of skin and other soft tissues are rare. Rebuilding a complete skeleton by comparing the size and morphology of bones to those of similar, better-known species is an inexact art, and reconstructing the muscles and other organs of the living animal is, at best, a process of educated guesswork.
The tallest and heaviest dinosaur known from good skeletons is Giraffatitan brancai
Giraffatitan
Giraffatitan, meaning "giraffe titan", is a genus of sauropod dinosaur that lived during the late Jurassic Period . It was originally named as an African species of Brachiosaurus...

(previously classified as a species of Brachiosaurus
Brachiosaurus
Brachiosaurus is a genus of sauropod dinosaur from the Jurassic Morrison Formation of North America. It was first described by Elmer S. Riggs in 1903 from fossils found in the Grand River Canyon of western Colorado, in the United States. Riggs named the dinosaur Brachiosaurus altithorax,...

). Its remains were discovered in Tanzania
Tanzania
The United Republic of Tanzania is a country in East Africa bordered by Kenya and Uganda to the north, Rwanda, Burundi, and the Democratic Republic of the Congo to the west, and Zambia, Malawi, and Mozambique to the south. The country's eastern borders lie on the Indian Ocean.Tanzania is a state...

 between 1907–12. Bones from several similar-sized individuals were incorporated into the skeleton now mounted and on display at the Museum für Naturkunde Berlin; this mount is 12 metres (39.4 ft) tall and 22.5 metres (73.8 ft) long, and would have belonged to an animal that weighed between and  kilograms ( and  lb). The longest complete dinosaur is the 27-meter (89 ft) long Diplodocus
Diplodocus
Diplodocus , or )is a genus of diplodocid sauropod dinosaur 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 "double" and "beam", in reference to its double-beamed chevron bones...

, which was discovered in Wyoming
Wyoming
Wyoming is a state in the mountain region of the Western United States. The western two thirds of the state is covered mostly with the mountain ranges and rangelands in the foothills of the Eastern Rocky Mountains, while the eastern third of the state is high elevation prairie known as the High...

 in the United States
United States
The United States of America is a federal constitutional republic comprising fifty states and a federal district...

 and displayed in Pittsburgh's
Pittsburgh, Pennsylvania
Pittsburgh is the second-largest city in the US Commonwealth of Pennsylvania and the county seat of Allegheny County. Regionally, it anchors the largest urban area of Appalachia and the Ohio River Valley, and nationally, it is the 22nd-largest urban area in the United States...

 Carnegie Natural History Museum in 1907.

There were larger dinosaurs, but knowledge of them is based entirely on a small number of fragmentary fossils. Most of the largest herbivorous
Herbivore
Herbivores are organisms that are anatomically and physiologically adapted to eat plant-based foods. Herbivory is a form of consumption in which an organism principally eats autotrophs such as plants, algae and photosynthesizing bacteria. More generally, organisms that feed on autotrophs in...

 specimens on record were all discovered in the 1970s or later, and include the massive Argentinosaurus
Argentinosaurus
Argentinosaurus is a genus of titanosaur sauropod dinosaur first discovered by Guillermo Heredia in Argentina. The generic name refers to the country in which it was discovered...

, which may have weighed to  kilograms (90 to 110 short tons); some of the longest were the 33.5 metres (109.9 ft) long Diplodocus hallorum
Diplodocus
Diplodocus , or )is a genus of diplodocid sauropod dinosaur 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 "double" and "beam", in reference to its double-beamed chevron bones...

(formerly Seismosaurus) and the 33 metres (108.3 ft) long Supersaurus
Supersaurus
Supersaurus is a genus of diplodocid sauropod dinosaur discovered by Vivian Jones of Delta, Colorado, in the Upper Jurassic Morrison Formation of Colorado in 1972. The fossil remains came from the Brushy Basin Member of the formation, dating to about 153 million years ago...

; and the tallest, the 18 metres (59.1 ft) tall Sauroposeidon
Sauroposeidon
Sauroposeidon is a genus of sauropod dinosaur known from four neck vertebrae that were found in the southeastern portion of the US state of Oklahoma. The fossils were found in rocks dating to the Early Cretaceous, a period when the sauropods of North America had diminished in both size and...

, which could have reached a sixth-floor window. The heaviest and longest of them all may have been Amphicoelias fragillimus
Amphicoelias
Amphicoelias is a genus of herbivorous sauropod dinosaur that includes what may be the largest dinosaur ever discovered, A. fragillimus. Based on surviving descriptions of a single fossil bone, A. fragillimus may have been the longest known vertebrate at in length, and may have had a mass of up...

, known only from a now lost partial vertebral neural arch described in 1878. Extrapolating from the illustration of this bone, the animal may have been 58 metres (190.3 ft) long and weighed over kg ( lb). The largest known carnivorous
Carnivore
A carnivore meaning 'meat eater' is an organism that derives its energy and nutrient requirements from a diet consisting mainly or exclusively of animal tissue, whether through predation or scavenging...

 dinosaur was Spinosaurus
Spinosaurus
Spinosaurus is a genus of theropod dinosaur which lived in what is now North Africa, from the lower Albian to lower Cenomanian stages of the Cretaceous period, about 112 to 97 million years ago. This genus was first known from Egyptian remains discovered in 1912 and described by German...

, reaching a length of 16 to 18 meters (50 to 60 ft), and weighing in at 8150 kg ( lb). Other large meat-eaters included Giganotosaurus
Giganotosaurus
Giganotosaurus is a genus of carcharodontosaurid dinosaur that lived around 97 million years ago during the early Cenomanian stage of the Late Cretaceous Period. It included some of the largest known terrestrial carnivores, slightly larger than the largest Tyrannosaurus, but smaller than the...

, Carcharodontosaurus
Carcharodontosaurus
Carcharodontosaurus was a gigantic carnivorous carcharodontosaurid dinosaur that lived around 100 to 93 million years ago, during the late Albian to early Cenomanian stages of the mid-Cretaceous Period...

and Tyrannosaurus
Tyrannosaurus
Tyrannosaurus meaning "tyrant," and sauros meaning "lizard") is a genus of coelurosaurian theropod dinosaur. The species Tyrannosaurus rex , commonly abbreviated to T. rex, is a fixture in popular culture. It lived throughout what is now western North America, with a much wider range than other...

.

Not including modern birds, the smallest known dinosaurs known were about the size of a pigeon. Not surprisingly, the smallest dinosaurs were theropods closely related to birds. Anchiornis
Anchiornis
Anchiornis is a genus of small, feathered, deinonychosaurian dinosaur. The genus Anchiornis contains the type species Anchiornis huxleyi, named in honor of Thomas Henry Huxley, an early proponent of biological evolution, and the first to propose a close evolutionary relationship between birds and...

and Epidexipteryx
Epidexipteryx
Epidexipteryx is a genus of small avialan dinosaur, known from one fossil specimen in the collection of the Institute of Vertebrate Paleontology and Paleoanthropology in Beijing. Epidexipteryx represents the earliest known example of ornamental feathers in the fossil record. The type specimen is...

both had a total skeletal length of under 35 centimeters (1.1 ft). Anchiornis is currently the smallest dinosaur described from an adult specimen, with an estimated weight of 110 grams. The smallest herbivorous dinosaurs included Microceratus and Wannanosaurus
Wannanosaurus
Wannanosaurus is a genus of basal pachycephalosaurian dinosaur from the Campanian-age Upper Cretaceous Xiaoyan Formation, about 80 million years ago in what is now Anhui, China...

, at about 60 cm (2 ft) long each.

Behavior


Interpretations of dinosaur behavior are generally based on the pose of body fossils and their habitat
Habitat (ecology)
A habitat is an ecological or environmental area that is inhabited by a particular species of animal, plant or other type of organism...

, computer simulation
Computer simulation
A computer simulation, a computer model, or a computational model is a computer program, or network of computers, that attempts to simulate an abstract model of a particular system...

s of their biomechanics
Biomechanics
Biomechanics is the application of mechanical principles to biological systems, such as humans, animals, plants, organs, and cells. Perhaps one of the best definitions was provided by Herbert Hatze in 1974: "Biomechanics is the study of the structure and function of biological systems by means of...

, and comparisons with modern animals in similar ecological niche
Ecological niche
In ecology, a niche is a term describing the relational position of a species or population in its ecosystem to each other; e.g. a dolphin could potentially be in another ecological niche from one that travels in a different pod if the members of these pods utilize significantly different food...

s. As such, the current understanding of dinosaur behavior relies on speculation, and will likely remain controversial for the foreseeable future. However, there is general agreement that some behaviors which are common in crocodiles and birds, dinosaurs' closest living relatives, were also common among dinosaurs.

The first potential evidence of herd
Herd
Herd refers to a social grouping of certain animals of the same species, either wild or domestic, and also to the form of collective animal behavior associated with this or as a verb, to herd, to its control by another species such as humans or dogs.The term herd is generally applied to mammals,...

ing behavior was the 1878 discovery of 31 Iguanodon
Iguanodon
Iguanodon is a genus of ornithopod dinosaur that lived roughly halfway between the first of the swift bipedal hypsilophodontids and the ornithopods' culmination in the duck-billed dinosaurs...

dinosaurs which were then thought to have perished together in Bernissart
Bernissart
Bernissart is a Walloon municipality located in the Belgian province of Hainaut. On January 1, 2006 Bernissart had a total population of 11,458. The total area is 43.42 km², which gives a population density of 264 inhabitants per km²....

, Belgium
Belgium
Belgium , officially the Kingdom of Belgium, is a federal state in Western Europe. It is a founding member of the European Union and hosts the EU's headquarters, and those of several other major international organisations such as NATO.Belgium is also a member of, or affiliated to, many...

, after they fell into a deep, flooded sinkhole
Sinkhole
A sinkhole, also known as a sink, shake hole, swallow hole, swallet, doline or cenote, is a natural depression or hole in the Earth's surface caused by karst processes — the chemical dissolution of carbonate rocks or suffosion processes for example in sandstone...

 and drowned. Other mass-death sites have been subsequently discovered. Those, along with multiple trackways, suggest that gregarious behavior was common in many dinosaur species. Trackways of hundreds or even thousands of herbivores indicate that duck-bills
Hadrosaurid
Hadrosaurids or duck-billed dinosaurs are members of the family Hadrosauridae, and include ornithopods such as Edmontosaurus and Parasaurolophus. They were common herbivores in the Upper Cretaceous Period of what are now Asia, Europe and North America. They are descendants of the Upper...

 (hadrosaurids) may have moved in great herds, like the American Bison
American Bison
The American bison , also commonly known as the American buffalo, is a North American species of bison that once roamed the grasslands of North America in massive herds...

 or the African Springbok
Springbok Antelope
The springbok is a medium-sized brown and white gazelle that stands about high. Springbok males weigh between and the females between...

. Sauropod tracks document that these animals traveled in groups composed of several different species, at least in Oxfordshire
Oxfordshire
Oxfordshire is a county in the South East region of England, bordering on Warwickshire and Northamptonshire , Buckinghamshire , Berkshire , Wiltshire and Gloucestershire ....

, England, although there is not evidence for specific herd structures. Dinosaurs may have congregated in herds for defense, for migratory
Bird migration
Bird migration is the regular seasonal journey undertaken by many species of birds. Bird movements include those made in response to changes in food availability, habitat or weather. Sometimes, journeys are not termed "true migration" because they are irregular or in only one direction...

 purposes, or to provide protection for their young. There is evidence that many types of dinosaurs, including various theropods, sauropods, ankylosaurians, ornithopods, and ceratopsians, formed aggregations of immature individuals. One example is a site in Inner Mongolia
Inner Mongolia
Inner Mongolia is an autonomous region of the People's Republic of China, located in the northern region of the country. Inner Mongolia shares an international border with the countries of Mongolia and the Russian Federation...

 that has yielded the remains of over 20 Sinornithomimus
Sinornithomimus
Sinornithomimus is a genus of ornithomimid theropod dinosaur found in 1997, in the early Late Cretaceous strata of the Ulansuhai Formation located at Alshanzuo Banner, Nei Mongol Autonomous Region, Northern China.-Discovery and naming:...

, 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 hunter
Pack hunter
A pack hunter is a predator belonging to the animal kingdom, which has evolved to hunt its prey by working together with other members of its species. Normally, such animals are closely related. The most commonly known pack hunter is the Gray Wolf, the ancestor of all breeds of domesticated dogs...

s working together to bring down large prey. However, this lifestyle is uncommon among the modern relatives of dinosaurs (crocodile
Crocodile
A crocodile is any species belonging to the family Crocodylidae . The term can also be used more loosely to include all extant members of the order Crocodilia: i.e...

s and other reptiles, and bird
Bird
Birds are feathered, winged, bipedal, endothermic , egg-laying, vertebrate animals. Around 10,000 living species and 188 families makes them the most speciose class of tetrapod vertebrates. They inhabit ecosystems across the globe, from the Arctic to the Antarctic. Extant birds range in size from...

s – Harris's Hawk
Harris's Hawk
The Harris's Hawk or Harris Hawk formerly known as the Bay-winged Hawk or Dusky Hawk, is a medium-large bird of prey which breeds from the southwestern United States south to Chile and central Argentina...

 is a well-documented exception), and the taphonomic
Taphonomy
Taphonomy is the study of decaying organisms over time and how they become fossilized . The term taphonomy was introduced to paleontology in 1940 by Russian scientist Ivan Efremov to describe the study of the transition of remains, parts, or products of organisms, from the biosphere, to the...

 evidence suggesting pack hunting in such theropods as Deinonychus
Deinonychus
Deinonychus was a genus of carnivorous dromaeosaurid dinosaur. There is one described species, Deinonychus antirrhopus. This 3.4 meter long dinosaur lived during the early Cretaceous Period, about 115–108 million years ago . Fossils have been recovered from the U.S...

and Allosaurus
Allosaurus
Allosaurus is a genus of large theropod dinosaur that lived 155 to 150 million years ago during the late Jurassic period . The name Allosaurus means "different lizard". It is derived from the Greek /allos and /sauros...

can also be interpreted as the results of fatal disputes between feeding animals, as is seen in many modern diapsid
Diapsid
Diapsids are a group of reptiles that developed two holes in each side of their skulls, about 300 million years ago during the late Carboniferous period. Living diapsids are extremely diverse, and include all crocodiles, lizards, snakes, and tuatara...

 predators.

Jack Horner's
Jack Horner (paleontologist)
John "Jack" R. Horner is an American paleontologist who discovered and named Maiasaura, providing the first clear evidence that some dinosaurs cared for their young. He is one of the best-known paleontologists in the United States...

 1978 discovery of a Maiasaura
Maiasaura
Maiasaura is a large duck-billed dinosaur genus that lived in the area currently covered by the state of Montana in the Upper Cretaceous Period , about 74 million years ago....

("good mother dinosaur") nest
Nest
A nest is a place of refuge to hold an animal's eggs or provide a place to live or raise offspring. They are usually made of some organic material such as twigs, grass, and leaves; or may simply be a depression in the ground, or a hole in a tree, rock or building...

ing ground in Montana
Montana
Montana is a state in the Western United States. The western third of Montana contains numerous mountain ranges. Smaller, "island ranges" are found in the central third of the state, for a total of 77 named ranges of the Rocky Mountains. This geographical fact is reflected in the state's name,...

 demonstrated that parental care continued long after birth among the ornithopod
Ornithopod
Ornithopods or members of the clade Ornithopoda are a group of ornithischian dinosaurs 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, and dominated the North American...

s. There is also evidence that other Cretaceous-era dinosaurs, like Patagonia
Patagonia
Patagonia is a region located in Argentina and Chile, integrating the southernmost section of the Andes mountains to the southwest towards the Pacific ocean and from the east of the cordillera to the valleys it follows south through Colorado River towards Carmen de Patagones in the Atlantic Ocean...

n titanosaur
Titanosaur
Titanosaurs were a diverse group of sauropod dinosaurs, which included Saltasaurus and Isisaurus. It includes some of the heaviest creatures ever to walk the earth, such as Argentinosaurus and Paralititan — which some believe have weighed up to 100 tonnes...

ian sauropods (1997 discovery), also nested in large groups. The Mongolia
Mongolia
Mongolia is a landlocked country in East and Central Asia. It is bordered by Russia to the north and China to the south, east and west. Although Mongolia does not share a border with Kazakhstan, its western-most point is only from Kazakhstan's eastern tip. Ulan Bator, the capital and largest...

n oviraptorid
Oviraptoridae
Oviraptoridae is a group of bird-like, herbivorous and omnivorous maniraptoran dinosaurs. Oviraptorids are characterized by their toothless, parrot-like beaks and, in some cases, elaborate crests. They were generally small, measuring between one and two metres long in most cases, though some...

 Citipati
Citipati
Citipati is a genus of oviraptorid theropod dinosaur from the Late Cretaceous Period of what is now Mongolia . It is one of the best-known oviraptorids, thanks to a number of well-preserved skeletons, including several specimens found in brooding positions atop nests of eggs...

was discovered in a chicken
Chicken
The chicken is a domesticated fowl, a subspecies of the Red Junglefowl. As one of the most common and widespread domestic animals, and with a population of more than 24 billion in 2003, there are more chickens in the world than any other species of bird...

-like brooding
Avian incubation
Incubation refers to the process by which certain oviparous animals hatch their eggs, and to the development of the embryo within the egg. The most vital factor of incubation is the constant temperature required for its development over a specific period. Especially in domestic fowl, the act of...

 position in 1993, which may mean it was covered with an insulating layer of feathers that kept the eggs
Egg (biology)
An egg is an organic vessel in which an embryo first begins to develop. In most birds, reptiles, insects, molluscs, fish, and monotremes, an egg is the zygote, resulting from fertilization of the ovum, which is expelled from the body and permitted to develop outside the body until the developing...

 warm. Parental care is also implied by other finds. For example, the fossilized remains of a grouping of Psittacosaurus
Psittacosaurus
Psittacosaurus is a genus of psittacosaurid ceratopsian dinosaur from the Early Cretaceous Period of what is now Asia, about 130 to 100 million years ago. It is notable for being the most species-rich dinosaur genus...

has been found, consisting of one adult and 34 juveniles; in this case, the large number of juveniles may be due to communal nesting. Additionally, a dinosaur embryo (pertaining to the prosauropod Massospondylus
Massospondylus
Massospondylus and ) is a genus of prosauropod dinosaur from the early Jurassic Period . It was described by Sir Richard Owen in 1854 from remains found in South Africa, and is thus one of the first dinosaurs to have been named...

) was found without teeth, indicating that some parental care was required to feed the young dinosaur. Trackways have also confirmed parental behavior among ornithopods from the Isle of Skye in northwestern Scotland
Scotland
Scotland is a country that is part of the United Kingdom. Occupying the northern third of the island of Great Britain, it shares a border with England to the south and is bounded by the North Sea to the east, the Atlantic Ocean to the north and west, and the North Channel and Irish Sea to the...

. Nests and eggs have been found for most major groups of dinosaurs, and it appears likely that dinosaurs communicated with their young, in a manner similar to modern birds and crocodiles.


The crests and frills
Neck frill
Neck frill is the popular term for 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 Lizard...

 of some dinosaurs, like the marginocephalia
Marginocephalia
Marginocephalia is a clade of ornithischian dinosaurs that includes the thick-skulled pachycephalosaurids, and horned ceratopsians. They were all herbivores, walking on two or four legs, and are characterized by a bony ridge or frill the back of the skull...

ns, theropods and lambeosaurines, may have been too fragile to be used for active defense, and so they were likely used for sexual or aggressive displays, though little is known about dinosaur mating and territorialism
Territory (animal)
In ethology the term territory refers to any sociographical area that an animal of a particular species consistently defends against conspecifics...

. 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
Gobi Desert
The Gobi is a large desert region in Asia. It covers parts of northern and northwestern China, and of southern Mongolia. The desert basins of the Gobi are bounded by the Altai Mountains and the grasslands and steppes of Mongolia on the north, by the Hexi Corridor and Tibetan Plateau to the...

 in 1971. It included a Velociraptor
Velociraptor
Velociraptor is a genus of dromaeosaurid theropod dinosaur that existed approximately 75 to 71 million years ago during the later part of the Cretaceous Period. Two species are currently recognized, although others have been assigned in the past. The type species is V. mongoliensis; fossils...

attacking a Protoceratops
Protoceratops
Protoceratops is a genus of sheep-sized herbivorous ceratopsian dinosaur, from the Upper Cretaceous Period of what is now Mongolia. It was a member of the Protoceratopsidae, a group of early horned dinosaurs...

, providing evidence that dinosaurs did indeed attack each other. Additional evidence for attacking live prey is the partially healed tail of an Edmontosaurus
Edmontosaurus
Edmontosaurus is a genus of crestless hadrosaurid dinosaur. It contains two species: Edmontosaurus regalis and Edmontosaurus annectens. Fossils of E. regalis have been found in rocks of western North America that date from the late Campanian stage of the Cretaceous Period 73 million years ago,...

, a hadrosaurid dinosaur; the tail is damaged in such a way that shows the animal was bitten by a tyrannosaur but survived. Cannibalism
Cannibalism (zoology)
In zoology, cannibalism is the act of one individual of a species consuming all or part of another individual of the same species as food. Cannibalism is a common ecological interaction in the animal kingdom and has been recorded for more than 1500 species...

 amongst some species of dinosaurs was confirmed by tooth marks found in Madagascar in 2003, involving the theropod Majungasaurus
Majungasaurus
Majungasaurus is a genus of abelisaurid theropod dinosaur that lived in Madagascar from 70 to 65.5 million years ago, at the end of the Cretaceous Period. Only one species has been identified...

.

Comparisons between the scleral ring
Sclerotic ring
Sclerotic rings are rings of bone found in the eyes of several groups of vertebrate animals, except for mammals and crocodilians. They can be made up of single bones or small bones together. They are believed to have a role in supporting the eye, especially in animals whose eyes are not spherical,...

s 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
Dromaeosauridae
Dromaeosauridae is a family of bird-like theropod dinosaurs. They were small- to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from Greek dromeus meaning 'runner' and sauros meaning 'lizard'...

, Juravenator
Juravenator
Juravenator is a genus of small coelurosaurian dinosaur, which lived in the area which would someday become the Jura mountains of Germany, about 151 or 152 million years ago. It is known from a single, juvenile specimen...

, and Megapnosaurus
Megapnosaurus
Megapnosaurus was a dinosaur of the theropod family Coelophysidae, formerly called Syntarsus , living during the Early Jurassic. It was renamed by American entomologist Dr. Michael Ivie , Polish Australian Dr...

were likely nocturnal. Large and medium-sized herbivorous
Herbivore
Herbivores are organisms that are anatomically and physiologically adapted to eat plant-based foods. Herbivory is a form of consumption in which an organism principally eats autotrophs such as plants, algae and photosynthesizing bacteria. More generally, organisms that feed on autotrophs in...

 and omnivorous
Omnivore
Omnivores are species that eat both plants and animals as their primary food source...

 dinosaurs such as ceratopsia
Ceratopsia
Ceratopsia or Ceratopia is a group of herbivorous, beaked dinosaurs which thrived in what are now North America, Europe, and Asia, during the Cretaceous Period, although ancestral forms lived earlier, in the Jurassic. The earliest known ceratopsian, Yinlong downsi, lived between 161.2 and 155.7...

ns, sauropodomorphs
Sauropodomorpha
Sauropodomorpha is an extinct clade of long-necked, herbivorous, saurischian dinosaurs which includes the sauropods and their ancestral relatives. Sauropods generally grew to very large sizes, had long necks and tails, were quadrupedal, and became the largest animals to ever walk the Earth. The...

, hadrosaurids, ornithomimosaurs
Ornithomimosauria
The Ornithomimosauria, ornithomimosaurs or ostrich dinosaurs were theropod dinosaurs which bore a superficial resemblance to modern ostriches. They were fast, omnivorous or herbivorous dinosaurs from the Cretaceous Period of Laurasia...

 may have been cathemeral
Cathemeral
A cathemeral organism is one that has sporadic and random intervals of activity during the day or night in which food is acquired, socializing with other organisms occurs, and any other activities necessary for livelihood are performed...

, active during short intervals throughout the day, although the small ornithischia
Ornithischia
Ornithischia or Predentata is an extinct order of beaked, herbivorous dinosaurs. The name ornithischia is derived from the Greek ornitheos meaning 'of a bird' and ischion meaning 'hip joint'...

n Agilisaurus
Agilisaurus
Agilisaurus is a genus of ornithischian dinosaur from the Middle Jurassic Period of what is now eastern Asia. The name is derived from the Latin agilis meaning 'agile' and the Greek sauros meaning 'lizard', and refers to the agility suggested by its lightweight skeleton and long legs...

was inferred to be diurnal.

Based on current fossil evidence from dinosaurs such as Oryctodromeus
Oryctodromeus
Oryctodromeus was a genus of small ornithopod dinosaur. Fossils are known from the middle Cretaceous Blackleaf Formation of southwestern Montana and the Wayan Formation of southeastern Idaho, both of the Cenomanian stage, roughly 95 million years ago...

, some herbivorous species seem to have led a partially fossorial
Fossorial
A fossorial organism is one that is adapted to digging and life underground such as the badger, the naked mole rat, and the mole salamanders Ambystomatidae...

 (burrowing) lifestyle, and some bird-like species may have been arboreal (tree climbing), most notably primitive dromaeosaurids
Dromaeosauridae
Dromaeosauridae is a family of bird-like theropod dinosaurs. They were small- to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from Greek dromeus meaning 'runner' and sauros meaning 'lizard'...

 such as Microraptor
Microraptor
Microraptor is a genus of small, four-winged dromaeosaurid dinosaurs. Numerous well-preserved fossil specimens have been recovered from Liaoning, China...

and the enigmatic scansoriopterygids
Scansoriopterygidae
Scansoriopterygidae is a family of maniraptoran dinosaurs, known from three well-preserved fossils unearthed in the Daohugou fossil beds of Liaoning, China....

. However, most 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
Biomechanics
Biomechanics is the application of mechanical principles to biological systems, such as humans, animals, plants, organs, and cells. Perhaps one of the best definitions was provided by Herbert Hatze in 1974: "Biomechanics is the study of the structure and function of biological systems by means of...

, in particular, 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 diplodocid
Diplodocid
Diplodocids, or members of the family Diplodocidae , are a group of sauropod dinosaurs. The family includes some of the longest creatures ever to walk the earth, including Diplodocus and Supersaurus, which may have reached lengths of up to .-Description:While still massive, when compared to the...

s could create sonic boom
Sonic boom
A sonic boom is the sound associated with the shock waves created by an object traveling through the air faster than the speed of sound. Sonic booms generate enormous amounts of sound energy, sounding much like an explosion...

s via whip
Whip
A whip is a tool traditionally used by humans to exert control over animals or other people, through pain compliance or fear of pain, although in some activities whips can be used without use of pain, such as an additional pressure aid in dressage...

-like tail snapping, and whether sauropods could float.

Communication and vocalization


The nature of dinosaur communication
Animal communication
Animal communication is any behavior on the part of one animal that has an effect on the current or future behaviour of another animal. The study of animal communication, is sometimes called Zoosemiotics has played an important part in the...

 remains enigmatic, and is an active area of research. In 2008, paleontologist Phil Senter examined the evidence for vocalization in Mesozoic animal life, including dinosaurs. Senter found that, contrary to popular depictions of roaring dinosaurs in motion pictures, it is likely that most dinosaurs were not capable of creating any vocalizations
Animal communication
Animal communication is any behavior on the part of one animal that has an effect on the current or future behaviour of another animal. The study of animal communication, is sometimes called Zoosemiotics has played an important part in the...

. To draw this conclusion, Senter studied the distribution of vocal organs in reptiles and birds. He found that vocal cords in the larynx
Larynx
The larynx , commonly called the voice box, is an organ in the neck of amphibians, reptiles and mammals involved in breathing, sound production, and protecting the trachea against food aspiration. It manipulates pitch and volume...

 probably evolved multiple times among reptiles, including crocodilia
Crocodilia
Crocodilia is an order of large reptiles that appeared about 84 million years ago in the late Cretaceous Period . They are the closest living relatives of birds, as the two groups are the only known survivors of the Archosauria...

ns, which are able to produce guttural roars. Birds, on the other hand, lack a larynx. Instead, bird calls are produced by the syrinx
Syrinx (biology)
Syrinx is the name for the vocal organ of birds. Located at the base of a bird's trachea, it produces sounds without the vocal cords of mammals. The sound is produced by vibrations of some or all of the membrana tympaniformis and the pessulus caused by air flowing through the syrinx...

, a vocal organ found only in birds, and which is not related to the larynx, meaning it evolved independently from the vocal organs in reptiles. The syrinx depends on the air sac system in birds to function; specifically, it requires the presence of a clavicular air sac near the wishbone or collar bone. This air sac leaves distinctive marks or opening on the bones, including a distinct opening in the upper arm bone (humerus). While many dinosaurs show evidence of extensive air sac systems, almost none possess the clavicular air sac necessary to vocalize (one exception, Aerosteon
Aerosteon
Aerosteon is a genus of allosauroid theropod dinosaur from the Late Cretaceous period of Argentina. Its remains were discovered in 1996 in the Anacleto Formation, dating to the Santonian stage . The type and only known species is A. riocoloradensis...

, probably evolved its clavicular air sac independently of birds for reasons other than vocalization).

The most primitive animals with evidence of a vocalizing syrinx are the enantironithine birds
Enantiornithes
Enantiornithes is an extinct group of primitive birds. They were the most abundant and diverse avialans of the Mesozoic. Almost all retained teeth and clawed fingers on each wing, but otherwise looked much like modern birds externally. Over 50 species of Enantiornithines have been named, but some...

. Any bird-line archosaurs more primitive than this probably did not make vocal calls. Rather, several lines of evidence suggest that dinosaurs used primarily visual communication, in the form of distinctive-looking (and possibly brightly colored) horns, frills, crests, sails and feathers. This is similar to some modern reptile groups such as lizards, in which many forms are largely silent (though like dinosaurs they possess well-developed senses of hearing) but use complex coloration and display behaviors to communicate.

Also, though they may not have been able to vocalize, some dinosaurs may have used other methods of producing sound for communication. Modern animals, including reptiles and birds, use a wide variety of non-vocal sound communication, including hissing, jaw grinding or clapping, use of environment (such as splashing), and wing beating (which would have been possible in winged maniraptora
Maniraptora
Maniraptora is a clade of coelurosaurian dinosaurs which includes the birds and the dinosaurs that were more closely related to them than to Ornithomimus velox. It contains the major subgroups Avialae, Deinonychosauria, Oviraptorosauria and Therizinosauria. Ornitholestes and the Alvarezsauroidea...

n dinosaurs).

Some studies have suggested that the hollow crests of the lambeosaurines may have functioned as resonance chamber
Resonance chamber
A resonance chamber uses resonance to amplify sound. The chamber has interior surfaces which reflect an acoustic wave. When a wave enters the chamber, it bounces back and forth within the chamber with low loss...

s used for a wide range of vocalizations. However, Senter (2008) noted that such chambers are also used in modern non-vocal animals to accentuate or deepen non-vocal sounds like hissing. For example, many snakes, which lack vocal cords, have resonating chambers in the skull.

Reproductive biology


A discovery of features in a Tyrannosaurus rex
Tyrannosaurus
Tyrannosaurus meaning "tyrant," and sauros meaning "lizard") is a genus of coelurosaurian theropod dinosaur. The species Tyrannosaurus rex , commonly abbreviated to T. rex, is a fixture in popular culture. It lived throughout what is now western North America, with a much wider range than other...

skeleton
Skeleton
The skeleton is the body part that forms the supporting structure of an organism. There are two different skeletal types: the exoskeleton, which is the stable outer shell of an organism, and the endoskeleton, which forms the support structure inside the body.In a figurative sense, skeleton can...

 provided evidence of medullary bone in dinosaurs and, for the first time, allowed paleontologists to establish the sex of a dinosaur. When laying eggs, female birds grow a special type of bone between the hard outer bone and the marrow
Bone marrow
Bone marrow is the flexible tissue found in the interior of bones. In humans, bone marrow in large bones produces new blood cells. On average, bone marrow constitutes 4% of the total body mass of humans; in adults weighing 65 kg , bone marrow accounts for approximately 2.6 kg...

 of their limbs. This medullary bone, which is rich in calcium
Calcium
Calcium is the chemical element with the symbol Ca and atomic number 20. It has an atomic mass of 40.078 amu. Calcium is a soft gray alkaline earth metal, and is the fifth-most-abundant element by mass in the Earth's crust...

, is used to make eggshells. The presence of endosteally derived bone tissues lining the interior marrow cavities of portions of the Tyrannosaurus rex specimen's hind limb suggested that T. rex used similar reproductive strategies, and revealed the specimen to be female. Further research has found medullary bone in the theropod Allosaurus
Allosaurus
Allosaurus is a genus of large theropod dinosaur that lived 155 to 150 million years ago during the late Jurassic period . The name Allosaurus means "different lizard". It is derived from the Greek /allos and /sauros...

and the ornithopod Tenontosaurus
Tenontosaurus
Tenontosaurus is a genus of medium- to large-sized ornithopod dinosaur. The genus is known from the late Aptian to Albian ages of the middle Cretaceous period sediments of western North America, dating between 115 to 108 million years ago...

. 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 dinosaurs in general produced medullary tissue. Medullary bone has been found in specimens of sub-adult size, which suggests that dinosaurs reached sexual maturity rather quickly for such large animals.

Waste


Like other reptile
Reptile
Reptiles are members of a class of air-breathing, ectothermic vertebrates which are characterized by laying shelled eggs , and having skin covered in scales and/or scutes. They are tetrapods, either having four limbs or being descended from four-limbed ancestors...

s, dinosaurs are primarily uricotelic, that is, their kidney
Kidney
The kidneys, organs with several functions, serve essential regulatory roles in most animals, including vertebrates and some invertebrates. They are essential in the urinary system and also serve homeostatic functions such as the regulation of electrolytes, maintenance of acid–base balance, and...

s extract nitrogenous wastes from their bloodstream and excrete it as uric acid
Uric acid
Uric acid is a heterocyclic compound of carbon, nitrogen, oxygen, and hydrogen with the formula C5H4N4O3. It forms ions and salts known as urates and acid urates such as ammonium acid urate. Uric acid is created when the body breaks down purine nucleotides. High blood concentrations of uric acid...

 instead of urea
Urea
Urea or carbamide is an organic compound with the chemical formula CO2. The molecule has two —NH2 groups joined by a carbonyl functional group....

 or ammonia
Ammonia
Ammonia is a compound of nitrogen and hydrogen with the formula . It is a colourless gas with a characteristic pungent odour. Ammonia contributes significantly to the nutritional needs of terrestrial organisms by serving as a precursor to food and fertilizers. Ammonia, either directly or...

 via the ureters into the intestine. In most living species, uric acid is excreted along with feces as a semisolid waste. However, at least some modern birds (such as hummingbird
Hummingbird
Hummingbirds are birds that comprise the family Trochilidae. They are among the smallest of birds, most species measuring in the 7.5–13 cm range. Indeed, the smallest extant bird species is a hummingbird, the 5-cm Bee Hummingbird. They can hover in mid-air by rapidly flapping their wings...

s) can be facultatively ammonotelic, excreting most of the nitrogenous wastes as ammonia. They also excrete creatine
Creatine
Creatine is a nitrogenous organic acid that occurs naturally in vertebrates and helps to supply energy to all cells in the body, primarily muscle. This is achieved by increasing the formation of Adenosine triphosphate...

, rather than creatinine
Creatinine
Creatinine is a break-down product of creatine phosphate in muscle, and is usually produced at a fairly constant rate by the body...

 like mammals. This material, as well as the output of the intestines, emerges from the cloaca
Cloaca
In zoological anatomy, a cloaca is the posterior opening that serves as the only such opening for the intestinal, reproductive, and urinary tracts of certain animal species...

. In addition, many species regurgitate pellets
Pellet (ornithology)
A pellet, in ornithology, is the mass of undigested parts of a bird's food that some bird species occasionally regurgitate. The contents of a bird's pellet depend on its diet, but can include the exoskeletons of insects, indigestible plant matter, bones, fur, feathers, bills, claws, and teeth...

, and fossil pellets that may have come from dinosaurs are known from as long ago as the Cretaceous period.

Physiology




A vigorous debate on the subject of temperature regulation in dinosaurs has been ongoing since the 1960s. Originally, scientists broadly disagreed as to whether dinosaurs were capable of regulating their body temperatures at all. More recently, dinosaur endothermy
Warm-blooded
The term warm-blooded is a colloquial term to describe animal species which have a relatively higher blood temperature, and maintain thermal homeostasis primarily through internal metabolic processes...

 has become the consensus view, and debate has focused on the mechanisms of temperature regulation.

After dinosaurs were discovered, paleontologists first posited that they were ectotherm
Ectotherm
An ectotherm, from the Greek εκτός "outside" and θερμός "hot", refers to organisms that control body temperature through external means. As a result, organisms are dependent on environmental heat sources and have relatively low metabolic rates. For example, many reptiles regulate their body...

ic creatures: "terrible lizard
Lizard
Lizards are a widespread group of squamate reptiles, with nearly 3800 species, ranging across all continents except Antarctica as well as most oceanic island chains...

s" as their name suggests. This supposed cold-bloodedness was used to imply that dinosaurs were relatively slow, sluggish organisms, even though many modern reptiles are fast and light-footed despite relying on external sources of heat to regulate their body temperature. The idea of dinosaurs as ectothermic and sluggish remained a prevalent view until Robert T. "Bob" Bakker
Robert T. Bakker
Robert T. Bakker is an American paleontologist who helped reshape modern theories about dinosaurs, particularly by adding support to the theory that some dinosaurs were endothermic...

, an early proponent of dinosaur endothermy, published an influential paper on the topic in 1968.

Modern evidence indicates that dinosaurs thrived in cooler temperate climates, and that at least some dinosaur species must have regulated their body temperature by internal biological means (aided by the animals' bulk in large species). Evidence of endothermy
Warm-blooded
The term warm-blooded is a colloquial term to describe animal species which have a relatively higher blood temperature, and maintain thermal homeostasis primarily through internal metabolic processes...

 in dinosaurs includes the discovery of polar dinosaurs in Australia
Polar dinosaurs in Australia
The South Polar dinosaurs proliferated during the Early Cretaceous while the continent of Australia was still linked to Antarctica to form East Gondwana, a continent that had rifted from Africa and drifted southward. Much of this southern continent lay inside the Antarctic Circle, and the climate...

 and Antarctica (where they would have experienced a cold, dark six-month winter), the discovery of dinosaurs whose feathers may have provided regulatory insulation, and analysis of blood-vessel structures within dinosaur bone that are typical of endotherms. Scientific debate continues regarding the specific ways in which dinosaurs regulated their temperature.

Complicating the debate is the fact that warm-bloodedness can emerge based on more than one mechanism. Most discussions of dinosaur endothermy tend to compare them with average-sized birds or mammals, which expend energy to elevate body temperature above that of the environment. Small birds and mammals also possess insulation
Thermal insulation
Thermal insulation is the reduction of the effects of the various processes of heat transfer between objects in thermal contact or in range of radiative influence. Heat transfer is the transfer of thermal energy between objects of differing temperature...

, such as fat
Fat
Fats consist of a wide group of compounds that are generally soluble in organic solvents and generally insoluble in water. Chemically, fats are triglycerides, triesters of glycerol and any of several fatty acids. Fats may be either solid or liquid at room temperature, depending on their structure...

, fur
Fur
Fur is a synonym for hair, used more in reference to non-human animals, usually mammals; particularly those with extensives body hair coverage. The term is sometimes used to refer to the body hair of an animal as a complete coat, also known as the "pelage". Fur is also used to refer to animal...

, or feather
Feather
Feathers are one of the epidermal growths that form the distinctive outer covering, or plumage, on birds and some non-avian theropod dinosaurs. They are considered the most complex integumentary structures found in vertebrates, and indeed a premier example of a complex evolutionary novelty. They...

s, which slows down heat loss. However, large mammals, such as elephants, face a different problem because of their relatively small ratio of surface area to volume (Haldane's
J. B. S. Haldane
John Burdon Sanderson Haldane FRS , known as Jack , was a British-born geneticist and evolutionary biologist. A staunch Marxist, he was critical of Britain's role in the Suez Crisis, and chose to leave Oxford and moved to India and became an Indian citizen...

 principle). This ratio compares the volume of an animal with the area of its skin: as an animal gets bigger, its surface area increases more slowly than its volume. At a certain point, the amount of heat radiated away through the skin drops below the amount of heat produced inside the body, forcing animals to use additional methods to avoid overheating. In the case of elephants, they have little hair as adults, have large ears which increase their surface area, and have behavioral adaptations as well (such as using the trunk to spray water on themselves and mud-wallowing). These behaviors increase cooling through evaporation.

Large dinosaurs would presumably have had to deal with similar issues; their body size suggest they lost heat relatively slowly to the surrounding air, and so could have been what are called inertial homeotherms
Gigantothermy
Gigantothermy is a phenomenon with significance in biology and paleontology, whereby large, bulky ectothermic animals are more easily able to maintain a constant, relatively high body temperature than smaller animals by virtue of their smaller surface area to volume ratio...

, animals that are warmer than their environments through sheer size rather than through special adaptations like those of birds or mammals. However, so far this theory fails to account for the numerous dog- and goat-sized dinosaur species, or the young of larger species.

Modern computerized tomography (CT) scans of a dinosaur
Thescelosaurus
Thescelosaurus was a genus of small ornithopod dinosaur that appeared at the very end of the Late Cretaceous period in North America. It was a member of the last dinosaurian fauna before the Cretaceous-Tertiary extinction event around 65.5 million years ago...

's chest cavity (conducted in 2000) found the apparent remnants of a four-chambered heart, much like those found in today's mammals and birds. The idea is controversial within the scientific community, coming under fire for bad anatomical science or simply wishful thinking. The question of how this find reflects on metabolic rate and dinosaur internal anatomy may be moot, though, regardless of the object's identity: both modern crocodilia
Crocodilia
Crocodilia is an order of large reptiles that appeared about 84 million years ago in the late Cretaceous Period . They are the closest living relatives of birds, as the two groups are the only known survivors of the Archosauria...

ns and bird
Bird
Birds are feathered, winged, bipedal, endothermic , egg-laying, vertebrate animals. Around 10,000 living species and 188 families makes them the most speciose class of tetrapod vertebrates. They inhabit ecosystems across the globe, from the Arctic to the Antarctic. Extant birds range in size from...

s, the closest living relatives of dinosaurs, have four-chambered hearts (albeit modified in crocodilians), and so dinosaurs probably had them as well.

Soft tissue and DNA


One of the best examples of soft-tissue impressions in a fossil dinosaur was discovered in Petraroia, Italy
Italy
Italy , officially the Italian Republic languages]] under the European Charter for Regional or Minority Languages. In each of these, Italy's official name is as follows:;;;;;;;;), is a unitary parliamentary republic in South-Central Europe. To the north it borders France, Switzerland, Austria and...

. The discovery was reported in 1998, and described the specimen of a small, very young coelurosaur, Scipionyx
Scipionyx
Scipionyx is a genus of theropod dinosaur from the Early Cretaceous of Italy, around 113 million years ago. There has been only one skeleton discovered, which is notable for the preservation of soft tissue and internal organs. It is the fossil of a juvenile only half a metre long...

 samniticus
. The fossil includes portions of the intestines, colon, liver, muscles, and windpipe of this immature dinosaur.

In the March 2005 issue of Science
Science (journal)
Science is the academic journal of the American Association for the Advancement of Science and is one of the world's top scientific journals....

, the paleontologist Mary Higby Schweitzer
Mary Higby Schweitzer
Mary Higby Schweitzer is a paleontologist at North Carolina State University known for leading the groups which discovered the remains of blood cells in dinosaur fossils and later discovered soft tissue remains in the Tyrannosaurus rex specimen MOR 1125, as well as evidence that the specimen was a...

 and her team announced the discovery of flexible material resembling actual soft tissue inside a 68-million-year-old Tyrannosaurus rex
Tyrannosaurus
Tyrannosaurus meaning "tyrant," and sauros meaning "lizard") is a genus of coelurosaurian theropod dinosaur. The species Tyrannosaurus rex , commonly abbreviated to T. rex, is a fixture in popular culture. It lived throughout what is now western North America, with a much wider range than other...

leg bone
Bone
Bones are rigid organs that constitute part of the endoskeleton of vertebrates. They support, and protect the various organs of the body, produce red and white blood cells and store minerals. Bone tissue is a type of dense connective tissue...

 from the Hell Creek Formation
Hell Creek Formation
The Hell Creek Formation is an intensely-studied division of Upper Cretaceous to lower Paleocene rocks in North America, named for exposures studied along Hell Creek, near Jordan, Montana...

 in Montana
Montana
Montana is a state in the Western United States. The western third of Montana contains numerous mountain ranges. Smaller, "island ranges" are found in the central third of the state, for a total of 77 named ranges of the Rocky Mountains. This geographical fact is reflected in the state's name,...

. 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 vessel
Blood vessel
The blood vessels are the part of the circulatory system that transports blood throughout the body. There are three major types of blood vessels: the arteries, which carry the blood away from the heart; the capillaries, which enable the actual exchange of water and chemicals between the blood and...

s, 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; study and interpretation of the material is ongoing.

The successful extraction of ancient DNA from dinosaur fossils has been reported on two separate occasions, but, upon further inspection and peer review
Peer review
Peer review is a process of self-regulation by a profession or a process of evaluation involving qualified individuals within the relevant field. Peer review methods are employed to maintain standards, improve performance and provide credibility...

, neither of these reports could be confirmed. However, a functional visual peptide
Peptide
Peptides are short polymers of amino acid monomers linked by peptide bonds. They are distinguished from proteins on the basis of size, typically containing less than 50 monomer units. The shortest peptides are dipeptides, consisting of two amino acids joined by a single peptide bond...

 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 protein
Protein
Proteins are biochemical compounds consisting of one or more polypeptides typically folded into a globular or fibrous form, facilitating a biological function. A polypeptide is a single linear polymer chain of amino acids bonded together by peptide bonds between the carboxyl and amino groups of...

s, including hemoglobin, have putatively been detected in dinosaur fossils.

Feathers and the 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
Gerhard Heilmann
Gerhard Heilmann was a Danish artist and paleontologist who created artistic depictions of Archeopteryx, Proavis and other early bird relatives apart from writing The Origin of Birds, a pioneering and influential account of bird evolution...

 in the early 20th century, the theory of birds as dinosaur descendants was abandoned in favor of the idea of their being descendants of generalized thecodont
Thecodont
Thecodont , now considered an obsolete term, was formerly used to describe a diverse range of early archosaurs that first appeared in the Latest Permian and flourished until the end of the Triassic period...

s, with the key piece of evidence being the supposed lack of clavicle
Clavicle
In human anatomy, the clavicle or collar bone is a long bone of short length that serves as a strut between the scapula and the sternum. It is the only long bone in body that lies horizontally...

s in dinosaurs. However, as later discoveries showed, clavicles (or a single fused wishbone
Furcula
The ' is a forked bone found in birds, formed by the fusion of the two clavicles. In birds, its function is the strengthening of the thoracic skeleton to withstand the rigors of flight....

, which derived from separate clavicles) were not actually absent; they had been found as early as 1924 in Oviraptor, but misidentified as an interclavicle
Interclavicle
An interclavicle is a bone which, in most tetrapods, is located between the clavicles. Therian mammals are the only tetrapods which never have an interclavicle, although some members of other groups also lack one. Monotremes, although part of the mammalian class, do have interclavicles...

. In the 1970s, John Ostrom
John Ostrom
John H. Ostrom was an American paleontologist who revolutionized modern understanding of dinosaurs in the 1960s, when he demonstrated that dinosaurs are more like big non-flying birds than they are like lizards , an idea first proposed by Thomas Henry Huxley in the 1860s, but which had garnered...

 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
Yixian Formation
The Yixian Formation is a geological formation in Jinzhou, Liaoning, People's Republic of China, that spans 11 million years during the early Cretaceous period...

, 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 maniraptora
Maniraptora
Maniraptora is a clade of coelurosaurian dinosaurs which includes the birds and the dinosaurs that were more closely related to them than to Ornithomimus velox. It contains the major subgroups Avialae, Deinonychosauria, Oviraptorosauria and Therizinosauria. Ornitholestes and the Alvarezsauroidea...

n coelurosaurs. A minority of scientists, most notably Alan Feduccia
Alan Feduccia
Alan Feduccia is a paleornithologist, specializing in the origins and phylogeny of birds. He is now Professor Emeritus at the University of North Carolina. Feduccia's principal authored works include two books, The Age of Birds and The Origin and Evolution of Birds, and numerous papers in various...

 and Larry Martin
Larry Martin
Larry Martin is an American vertebrate paleontologist and curator of the Natural History Museum and Biodiversity Research Center at the University of Kansas. Among Martin's work is research on the Triassic reptile Longisquama and theropod dinosaur Caudipteryx...

, 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
Convergent evolution
Convergent evolution describes the acquisition of the same biological trait in unrelated lineages.The wing is a classic example of convergent evolution in action. Although their last common ancestor did not have wings, both birds and bats do, and are capable of powered flight. The wings are...

 with dinosaurs.

Feathers



Archaeopteryx was the first fossil found which revealed a potential connection between dinosaurs and birds. It is considered a transitional fossil
Transitional fossil
A transitional fossil is any fossilized remains of a lifeform that exhibits characteristics of two distinct taxonomic groups. A transitional fossil is the fossil of an organism near the branching point where major individual lineages diverge...

, in that it displays features of both groups. Brought to light just two years after Darwin's seminal The Origin of Species
The Origin of Species
Charles Darwin's On the Origin of Species, published on 24 November 1859, is a work of scientific literature which is considered to be the foundation of evolutionary biology. Its full title was On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the...

, its discovery spurred the nascent debate between proponents of evolutionary biology and creationism
Creationism
Creationism is the religious beliefthat humanity, life, the Earth, and the universe are the creation of a supernatural being, most often referring to the Abrahamic god. As science developed from the 18th century onwards, various views developed which aimed to reconcile science with the Genesis...

. This early bird is so dinosaur-like that, without a clear impression of feathers in the surrounding rock, at least one specimen was mistaken for Compsognathus
Compsognathus
Compsognathus was a small, bipedal, carnivorous theropod dinosaur. The animal was the size of a turkey and lived around 150 million years ago, the early Tithonian stage of the late Jurassic Period, in what is now Europe. Paleontologists have found two well-preserved fossils, one in Germany...

.

Since the 1990s, a number of additional feathered dinosaurs
Feathered dinosaurs
The realization that dinosaurs are closely related to birds raised the obvious possibility of feathered dinosaurs. Fossils of Archaeopteryx include well-preserved feathers, but it was not until the early 1990s that clearly non-avialan dinosaur fossils were discovered with preserved feathers...

 have been found, providing even stronger evidence of the close relationship between dinosaurs and modern birds. Most of these specimens were unearthed in the lagerstätte of the Yixian Formation, Liaoning
Liaoning
' is a province of the People's Republic of China, located in the northeast of the country. Its one-character abbreviation is "辽" , a name taken from the Liao River that flows through the province. "Níng" means "peace"...

, northeastern China
China
Chinese civilization may refer to:* China for more general discussion of the country.* Chinese culture* Greater China, the transnational community of ethnic Chinese.* History of China* Sinosphere, the area historically affected by Chinese culture...

, which was part of an island continent during the Cretaceous. Though feathers have been found in only a few locations, it is possible that non-avian dinosaurs elsewhere in the world were also feathered. The lack of widespread fossil evidence for feathered non-avian dinosaurs may be because delicate features like skin and feathers are not often preserved by fossilization and thus are absent from the fossil record. To this point, protofeathers (thin, filament-like structures) are known from dinosaurs at the base of Coelurosauria, such as compsognathids
Compsognathidae
Compsognathidae is a family of small carnivorous dinosaurs, generally conservative in form, from the Jurassic and Cretaceous Periods. Compsognathids lie at or near the origin of feathers—skin impressions are known from four genera, Compsognathus, Sinosauropteryx, Sinocalliopteryx, and Juravenator...

 like Sinosauropteryx
Sinosauropteryx
Sinosauropteryx was the first genus of dinosaur outside of Avialae to be found with evidence of feathers. They were covered with "furry" coats of very simple filament-like feathers...

and tyrannosauroids
Tyrannosauroidea
Tyrannosauroidea is a superfamily of coelurosaurian theropod dinosaurs that includes the family Tyrannosauridae as well as more basal relatives. Tyrannosauroids lived on the Laurasian supercontinent beginning in the Jurassic Period...

 (Dilong
Dilong (dinosaur)
Dilong is a genus of small tyrannosauroid dinosaur. The only species is Dilong paradoxus. It is from the Lower Cretaceous Yixian Formation near Lujiatun, Beipiao, in the western Liaoning province of China. It lived about 130 million years ago...

), but barbed feathers are known only among the coelurosaur subgroup Maniraptora, which includes oviraptorosaurs, troodontids, dromaeosaurids, and birds. 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 protofeathers are the result of the decomposition of collagenous fiber that underlaid the dinosaurs' integument, and that maniraptoran dinosaurs with barbed feathers were not actually dinosaurs, but convergent
Convergent evolution
Convergent evolution describes the acquisition of the same biological trait in unrelated lineages.The wing is a classic example of convergent evolution in action. Although their last common ancestor did not have wings, both birds and bats do, and are capable of powered flight. The wings are...

 with dinosaurs. However, their views have for the most part not been accepted by other researchers, to the point that the question of the scientific nature of Feduccia's proposals has been raised.

Skeleton


Because feathers are often associated with birds, feathered dinosaurs are often touted as the missing link
Transitional fossil
A transitional fossil is any fossilized remains of a lifeform that exhibits characteristics of two distinct taxonomic groups. A transitional fossil is the fossil of an organism near the branching point where major individual lineages diverge...

 between birds and dinosaurs. However, the multiple skeletal features also shared by the two groups represent another important line of evidence for paleontologists
Paleontology
Paleontology "old, ancient", ὄν, ὀντ- "being, creature", and λόγος "speech, thought") is the study of prehistoric life. It includes the study of fossils to determine organisms' evolution and interactions with each other and their environments...

. Areas of the skeleton with important similarities include the neck, pubis
Pubis (bone)
In vertebrates, the pubic bone is the ventral and anterior of the three principal bones composing either half of the pelvis.It is covered by a layer of fat, which is covered by the mons pubis....

, wrist
Wrist
In human anatomy, the wrist is variously defined as 1) the carpus or carpal bones, the complex of eight bones forming the proximal skeletal segment of the hand;...

 (semi-lunate carpal), arm and pectoral girdle, furcula (wishbone), and breast bone
Keel (bird)
A keel or carina in bird anatomy is an extension of the sternum which runs axially along the midline of the sternum and extends outward, perpendicular to the plane of the ribs. The keel provides an anchor to which a bird's wing muscles attach, thereby providing adequate leverage for flight...

. Comparison of bird and dinosaur skeletons through cladistic analysis
Cladistics
Cladistics is a method of classifying species of organisms into groups called clades, which consist of an ancestor organism and all its descendants . For example, birds, dinosaurs, crocodiles, and all descendants of their most recent common ancestor form a clade...

 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 an investigation which was led by Patrick O'Connor of Ohio University
Ohio University
Ohio University is a public university located in the Midwestern United States in Athens, Ohio, situated on an campus...

. The lungs of theropod dinosaurs (carnivores that walked on two legs and had bird-like feet) likely pumped air into hollow sacs in their skeleton
Skeleton
The skeleton is the body part that forms the supporting structure of an organism. There are two different skeletal types: the exoskeleton, which is the stable outer shell of an organism, and the endoskeleton, which forms the support structure inside the body.In a figurative sense, skeleton can...

s, 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 a 2008 paper published in the online journal PLoS ONE
PLoS ONE
PLoS ONE is an open access peer-reviewed scientific journal published by the Public Library of Science since 2006. It covers primary research from any discipline within science and medicine. All submissions go through an internal and external pre-publication peer review but are not excluded on the...

, scientists described Aerosteon riocoloradensis
Aerosteon
Aerosteon is a genus of allosauroid theropod dinosaur from the Late Cretaceous period of Argentina. Its remains were discovered in 1996 in the Anacleto Formation, dating to the Santonian stage . The type and only known species is A. riocoloradensis...

, the skeleton of which supplies the strongest evidence to date of a dinosaur with a bird-like breathing system. CT-scanning
Computed tomography
X-ray computed tomography or Computer tomography , is a medical imaging method employing tomography created by computer processing...

 revealed the evidence of air sacs within the body cavity of the Aerosteon skeleton.

Behavioral evidence


Fossils of the troodont
Troodontidae
Troodontidae is a family of bird-like theropod dinosaurs. During most of the 20th century, troodontid fossils were few and scrappy and they have therefore been allied, at various times, with many dinosaurian lineages...

s Mei
Mei (dinosaur)
Mei is a genus of duck-sized troodontid dinosaur first unearthed by paleontologists in Liaoning, China in 2004. Mei lived during the Early Cretaceous Period. Its binomial name, Mei long means sleeping dragon...

and Sinornithoides
Sinornithoides
Sinornithoides is a genus of troodontid theropod dinosaur. It lived during the Early Cretaceous . It measured approximately 1 meter long...

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 deinonychosaur
Deinonychosauria
The Deinonychosauria were a clade of coelurosaurian theropod dinosaurs from the Late Jurassic and Cretaceous periods. These omnivores and carnivores are known for their switchblade-like second toe claws and for displaying numerous bird-like characteristics. The clade has been divided into two...

 and oviraptorosaur
Oviraptorosauria
Oviraptorosaurs are a group of feathered maniraptoran dinosaurs from the Cretaceous Period of what are now Asia and North America. They are distinct for their characteristically short, beaked, parrot - like skulls, with or without bony crests atop the head...

 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
Gizzard
The gizzard, also referred to as the ventriculus, gastric mill, and gigerium, is an organ found in the digestive tract of some animals, including birds, reptiles, earthworms and some fish. This specialized stomach constructed of thick, muscular walls is used for grinding up food; often rocks are...

 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 gastrolith
Gastrolith
A gastrolith, also called a stomach stone or gizzard stones, is a rock held inside a gastrointestinal tract. Gastroliths are retained in the muscular gizzard and used to grind food in animals lacking suitable grinding teeth. The grain size depends upon the size of the animal and the gastrolith's...

s.

Extinction of major groups


The discovery that birds are a type of dinosaur showed that dinosaurs in general are not, in fact, extinct as is commonly stated. However, all non-avian dinosaurs as well as many groups of birds did suddenly become extinct approximately 65 million years ago. Many other groups of animals also became extinct at this time, including ammonite
Ammonite
Ammonite, as a zoological or paleontological term, refers to any member of the Ammonoidea an extinct subclass within the Molluscan class Cephalopoda which are more closely related to living coleoids Ammonite, as a zoological or paleontological term, refers to any member of the Ammonoidea an extinct...

s (nautilus
Nautilus
Nautilus is the common name of marine creatures of cephalopod family Nautilidae, the sole extant family of the superfamily Nautilaceae and of its smaller but near equal suborder, Nautilina. It comprises six living species in two genera, the type of which is the genus Nautilus...

-like mollusks), mosasaur
Mosasaur
Mosasaurs are large extinct marine lizards. The first fossil remains were discovered in a limestone quarry at Maastricht on the Meuse in 1764...

s, plesiosaur
Plesiosaur
Plesiosauroidea is an extinct clade of carnivorous plesiosaur marine reptiles. Plesiosauroids, are known from the Jurassic and Cretaceous Periods...

s, pterosaur
Pterosaur
Pterosaurs were flying reptiles of the clade or order Pterosauria. They existed from the late Triassic to the end of the Cretaceous Period . Pterosaurs are the earliest vertebrates known to have evolved powered flight...

s, and many groups of mammal
Mammal
Mammals are members of a class of air-breathing vertebrate animals characterised by the possession of endothermy, hair, three middle ear bones, and mammary glands functional in mothers with young...

s. This mass extinction is known as the Cretaceous–Paleogene extinction event. The nature of the event that caused this mass extinction has been extensively studied since the 1970s; at present, several related theories are supported by paleontologists. Though the consensus is that an impact event was the primary cause of dinosaur extinction, some scientists cite other possible causes, or support the idea that a confluence of several factors was responsible for the sudden disappearance of dinosaurs from the fossil record.

At the peak of the Mesozoic, there were no polar ice cap
Polar ice cap
A polar ice cap is a high latitude region of a planet or natural satellite that is covered in ice. There are no requirements with respect to size or composition for a body of ice to be termed a polar ice cap, nor any geological requirement for it to be over land; only that it must be a body of...

s, and sea levels are estimated to have been from 100 to 250 meters (300 to 800 ft) higher than they are today. The planet's temperature was also much more uniform, with only 25 °C (45 °F) separating average polar temperatures from those at the equator. On average, atmospheric temperatures were also much higher; the poles, for example, were 50 °C (90 °F) warmer than today.

The atmosphere's composition during the Mesozoic was vastly different as well. Carbon dioxide levels were up to 12 times higher than today's levels, and oxygen formed 32 to 35% of the atmosphere, as compared to 21% today. However, by the late Cretaceous, the environment was changing dramatically. Volcanic activity was decreasing, which led to a cooling trend as levels of atmospheric carbon dioxide dropped. Oxygen levels in the atmosphere also started to fluctuate and would ultimately fall considerably. Some scientists hypothesize that climate change, combined with lower oxygen levels, might have led directly to the demise of many species. If the dinosaurs had respiratory systems similar to those commonly found in modern birds, it may have been particularly difficult for them to cope with reduced respiratory efficiency, given the enormous oxygen demands of their very large bodies.

Impact event




The asteroid collision theory, which was brought to wide attention in 1980 by Walter Alvarez
Walter Alvarez
Walter Alvarez is a professor in the Earth and Planetary Science department at the University of California, Berkeley. He is most widely known for the theory that dinosaurs were killed by an asteroid impact, developed in collaboration with his father, Nobel Prize winning physicist Luis...

 and colleagues, links the extinction event
Extinction event
An extinction event is a sharp decrease in the diversity and abundance of macroscopic life. They occur when the rate of extinction increases with respect to the rate of speciation...

 at the end of the Cretaceous period to a bolide impact approximately 65.5 million years ago. Alvarez et al. proposed that a sudden increase in iridium
Iridium
Iridium is the chemical element with atomic number 77, and is represented by the symbol Ir. A very hard, brittle, silvery-white transition metal of the platinum family, iridium is the second-densest element and is the most corrosion-resistant metal, even at temperatures as high as 2000 °C...

 levels, recorded around the world in the period's rock stratum, was direct evidence of the impact. The bulk of the evidence now suggests that a bolide 5 to 15 kilometers (3 to 9 mi) wide hit in the vicinity of the Yucatán Peninsula
Yucatán Peninsula
The Yucatán Peninsula, in southeastern Mexico, separates the Caribbean Sea from the Gulf of Mexico, with the northern coastline on the Yucatán Channel...

 (in southeastern Mexico
Mexico
The United Mexican States , commonly known as Mexico , is a federal constitutional republic in North America. It is bordered on the north by the United States; on the south and west by the Pacific Ocean; on the southeast by Guatemala, Belize, and the Caribbean Sea; and on the east by the Gulf of...

), creating the approximately 180 km (111.8 mi) Chicxulub Crater
Chicxulub Crater
The Chicxulub crater is an ancient impact crater buried underneath the Yucatán Peninsula in Mexico. Its center is located near the town of Chicxulub, after which the crater is named...

 and triggering the mass extinction. Scientists are not certain whether dinosaurs were thriving or declining before the impact event
Impact event
An impact event is the collision of a large meteorite, asteroid, comet, or other celestial object with the Earth or another planet. Throughout recorded history, hundreds of minor impact events have been reported, with some occurrences causing deaths, injuries, property damage or other significant...

. Some scientists propose that the meteorite caused a long and unnatural drop in Earth's atmospheric temperature, while others claim that it would have instead created an unusual heat wave. The consensus among scientists who support this theory is that the impact caused extinctions both directly (by heat from the meteorite impact) and also indirectly (via a worldwide cooling brought about when matter ejected from the impact crater reflected thermal radiation from the sun). Although the speed of extinction cannot be deduced from the fossil record alone, various models suggest that the extinction was extremely rapid, being down to hours rather than years.

In September 2007, U.S. researchers led by William Bottke of the Southwest Research Institute
Southwest Research Institute
Southwest Research Institute , headquartered in San Antonio, Texas, is one of the oldest and largest independent, nonprofit, applied research and development organizations in the United States...

 in Boulder, Colorado
Boulder, Colorado
Boulder is the county seat and most populous city of Boulder County and the 11th most populous city in the U.S. state of Colorado. Boulder is located at the base of the foothills of the Rocky Mountains at an elevation of...

, and Czech
Czech Republic
The Czech Republic is a landlocked country in Central Europe. The country is bordered by Poland to the northeast, Slovakia to the east, Austria to the south, and Germany to the west and northwest....

 scientists used computer
Computer
A computer is a programmable machine designed to sequentially and automatically carry out a sequence of arithmetic or logical operations. The particular sequence of operations can be changed readily, allowing the computer to solve more than one kind of problem...

 simulations to identify the probable source of the Chicxulub impact. They calculated a 90% probability that a giant asteroid named Baptistina, approximately 160 km (99.4 mi) in diameter, orbiting in the asteroid belt which lies between Mars
Mars
Mars is the fourth planet from the Sun in the Solar System. The planet is named after the Roman god of war, Mars. It is often described as the "Red Planet", as the iron oxide prevalent on its surface gives it a reddish appearance...

 and Jupiter
Jupiter
Jupiter is the fifth planet from the Sun and the largest planet within the Solar System. It is a gas giant with mass one-thousandth that of the Sun but is two and a half times the mass of all the other planets in our Solar System combined. Jupiter is classified as a gas giant along with Saturn,...

, was struck by a smaller unnamed asteroid about 55 km (35 mi) in diameter about 160 million years ago. The impact shattered Baptistina, creating a cluster which still exists today as the Baptistina family
Baptistina family
The Baptistina family is an asteroid family that was likely produced by the breakup of an asteroid across 80 million years ago following an impact with a smaller body. The largest presumed remnant of this parent asteroid is 298 Baptistina....

. Calculations indicate that some of the fragments were sent hurtling into earth-crossing orbits, one of which was the 10 km (6.2 mi) wide meteorite
Meteorite
A meteorite is a natural object originating in outer space that survives impact with the Earth's surface. Meteorites can be big or small. Most meteorites derive from small astronomical objects called meteoroids, but they are also sometimes produced by impacts of asteroids...

 which struck Mexico
Mexico
The United Mexican States , commonly known as Mexico , is a federal constitutional republic in North America. It is bordered on the north by the United States; on the south and west by the Pacific Ocean; on the southeast by Guatemala, Belize, and the Caribbean Sea; and on the east by the Gulf of...

's Yucatan
Yucatán
Yucatán officially Estado Libre y Soberano de Yucatán is one of the 31 states which, with the Federal District, comprise the 32 Federal Entities of Mexico. It is divided in 106 municipalities and its capital city is Mérida....

 peninsula
Peninsula
A peninsula is a piece of land that is bordered by water on three sides but connected to mainland. In many Germanic and Celtic languages and also in Baltic, Slavic and Hungarian, peninsulas are called "half-islands"....

 65 million years ago, creating the Chicxulub crater. In 2011, new data from the Wide-field Infrared Survey Explorer
Wide-field Infrared Survey Explorer
Wide-field Infrared Survey Explorer is a NASA infrared-wavelength astronomical space telescope launched on December 14, 2009, and decommissioned/hibernated on February 17, 2011 when its transmitter was turned off...

 revised the date of the collision which created the Baptistina family
Baptistina family
The Baptistina family is an asteroid family that was likely produced by the breakup of an asteroid across 80 million years ago following an impact with a smaller body. The largest presumed remnant of this parent asteroid is 298 Baptistina....

 to about 80 million years ago. This makes an asteroid from this family highly improbable to be the asteroid that created the Chicxulub Crater, as typically the process of resonance and collision of an asteroid takes many tens of millions of years.

A similar but more controversial explanation proposes that "passages of the [hypothetical] solar companion star Nemesis
Nemesis (star)
Nemesis is a hypothetical hard-to-detect red dwarf star, white dwarf star or brown dwarf, originally postulated in 1984 to be orbiting the Sun at a distance of about 95,000 AU , somewhat beyond the Oort cloud, to explain a perceived cycle of mass extinctions in the geological record, which seem to...

 through the Oort comet cloud
Oort cloud
The Oort cloud , or the Öpik–Oort cloud , is a hypothesized spherical cloud of comets which may lie roughly 50,000 AU, or nearly a light-year, from the Sun. This places the cloud at nearly a quarter of the distance to Proxima Centauri, the nearest star to the Sun...

 would trigger comet showers." One or more of these comets then collided with the Earth at approximately the same time, causing the worldwide extinction. As with the impact of a single asteroid, the end result of this comet bombardment would have been a sudden drop in global temperatures, followed by a protracted cool period.

Deccan Traps


Before 2000, arguments that the Deccan Traps
Deccan Traps
The Deccan Traps are a large igneous province located on the Deccan Plateau of west-central India and one of the largest volcanic features on Earth. They consist of multiple layers of solidified flood basalt that together are more than thick and cover an area of and a volume of...

 flood basalt
Flood basalt
A flood basalt or trap basalt is the result of a giant volcanic eruption or series of eruptions that coats large stretches of land or the ocean floor with basalt lava. Flood basalts have occurred on continental scales in prehistory, creating great plateaus and mountain ranges...

s caused the extinction
Extinction
In biology and ecology, extinction is the end of an organism or of a group of organisms , normally 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 recover may have been lost before this point...

 were usually linked to the view that the extinction was gradual, as the flood basalt events were thought to have started around 68 million years ago and lasted for over 2 million years. However, there is evidence that two thirds of the Deccan Traps were created in only 1 million years about 65.5 million years ago, and so these eruptions would have caused a fairly rapid extinction, possibly over a period of thousands of years, but still longer than would be expected from a single impact event.

The Deccan Traps could have caused extinction through several mechanisms, including the release into the air of dust and sulphuric aerosols, which might have blocked sunlight and thereby reduced photosynthesis in plants. In addition, Deccan Trap volcanism might have resulted in carbon dioxide emissions, which would have increased the greenhouse effect
Greenhouse effect
The greenhouse effect is a process by which thermal radiation from a planetary surface is absorbed by atmospheric greenhouse gases, and is re-radiated in all directions. Since part of this re-radiation is back towards the surface, energy is transferred to the surface and the lower atmosphere...

 when the dust and aerosols cleared from the atmosphere. Before the mass extinction of the dinosaurs, the release of volcanic gas
Volcanic gas
|250px|thumb|right|Image of the [[rhyolitic]] [[lava dome]] of [[Chaitén Volcano]] during its 2008-2010 eruption.Volcanic gases include a variety of substances given off by active volcanoes...

es during the formation of the Deccan Traps
Deccan Traps
The Deccan Traps are a large igneous province located on the Deccan Plateau of west-central India and one of the largest volcanic features on Earth. They consist of multiple layers of solidified flood basalt that together are more than thick and cover an area of and a volume of...

 "contributed to an apparently massive global warming. Some data point to an average rise in temperature of 8 °C (14 °F) in the last half million years before the impact [at Chicxulub
Chicxulub Crater
The Chicxulub crater is an ancient impact crater buried underneath the Yucatán Peninsula in Mexico. Its center is located near the town of Chicxulub, after which the crater is named...

]."

In the years when the Deccan Traps theory was linked to a slower extinction, Luis Alvarez
Luis Alvarez
Luis W. Alvarez was an American experimental physicist and inventor, who spent nearly all of his long professional career on the faculty of the University of California, Berkeley...

 (who died in 1988) replied that paleontologists were being misled by sparse data. While his assertion was not initially well-received, later intensive field studies of fossil beds lent weight to his claim. Eventually, most paleontologists began to accept the idea that the mass extinctions at the end of the Cretaceous were largely or at least partly due to a massive Earth impact. However, even Walter Alvarez has acknowledged that there were other major changes on Earth even before the impact, such as a drop in sea level
Sea level
Mean sea level is a measure of the average height of the ocean's surface ; used as a standard in reckoning land elevation...

 and massive volcanic eruptions that produced the Indian Deccan Traps, and these may have contributed to the extinctions.

Failure to adapt to changing conditions


Lloyd et al. (2008) noted that, in the Mid Cretaceous, the flowering, angiosperm plants became a major part of terrestrial ecosystem
Ecosystem
An ecosystem is a biological environment consisting of all the organisms living in a particular area, as well as all the nonliving , physical components of the environment with which the organisms interact, such as air, soil, water and sunlight....

s, which had previously been dominated by gymnosperm
Gymnosperm
The gymnosperms are a group of seed-bearing plants that includes conifers, cycads, Ginkgo, and Gnetales. The term "gymnosperm" comes from the Greek word gymnospermos , meaning "naked seeds", after the unenclosed condition of their seeds...

s such as conifers. Dinosaur coprolite
Coprolite
A coprolite is fossilized animal dung. Coprolites are classified as trace fossils as opposed to body fossils, as they give evidence for the animal's behaviour rather than morphology. The name is derived from the Greek words κοπρος / kopros meaning 'dung' and λιθος / lithos meaning 'stone'. They...

–fossilized dung–indicate that, while some ate angiosperms, most herbivorous dinosaurs ate mainly gymnosperms. Statistical analysis by Lloyd et al. concluded that, contrary to earlier studies, dinosaurs did not diversify very much in the Late Cretaceous. Lloyd et al. suggested that dinosaurs' failure to diversify as ecosystems were changing doomed them to extinction.

Possible Paleocene survivors



Non-avian dinosaur remains are occasionally found above the K–T boundary
K–T boundary
The K–T boundary is a geological signature, usually a thin band, dated to 65.5 ± 0.3 Ma ago. K is the traditional abbreviation for the Cretaceous period, and T is the abbreviation for the Tertiary period...

. In 2001, paleontologists Zielinski and Budahn reported the discovery of a single hadrosaur leg-bone fossil in the San Juan Basin, New Mexico, and described it as evidence of Paleocene dinosaurs
Paleocene dinosaurs
Paleocene dinosaurs describe families or genera of non-avian dinosaurs that may have survived the Cretaceous–Tertiary extinction event 65.5 million years ago...

. The formation in which the bone was discovered has been dated to the early Paleocene
Paleocene
The Paleocene or Palaeocene, the "early recent", is a geologic epoch that lasted from about . It is the first epoch of the Palaeogene Period in the modern Cenozoic Era...

 epoch, approximately 64.5 million years ago. If the bone was not re-deposited into that stratum
Stratum
In geology and related fields, a stratum is a layer of sedimentary rock or soil with internally consistent characteristics that distinguish it from other layers...

 by weathering action, it would provide evidence that some dinosaur populations may have survived at least a half million years into the Cenozoic Era. Other evidence includes the finding of dinosaur remains in the Hell Creek Formation
Hell Creek Formation
The Hell Creek Formation is an intensely-studied division of Upper Cretaceous to lower Paleocene rocks in North America, named for exposures studied along Hell Creek, near Jordan, Montana...

 up to 1.3 meters (51 in) above ( years later than) the K–T boundary. Similar reports have come from other parts of the world, including China. Many scientists, however, dismissed the supposed Paleocene dinosaurs as re-worked, that is, washed out of their original locations and then re-buried in much later sediments. However, direct dating of the bones themselves has supported the later date, with U–Pb dating methods resulting in a precise age of 64.8 ± 0.9 million years ago. If correct, the presence of a handful of dinosaurs in the early Paleocene would not change the underlying facts of the extinction.

History of discovery


Dinosaur fossils have been known for millennia, although their true nature was not recognized. The Chinese, whose modern word for dinosaur is konglong (恐龍, or "terrible dragon"), considered them to be dragon
Chinese dragon
Chinese dragons are legendary creatures in Chinese mythology and folklore, with mythic counterparts among Japanese, Korean, Vietnamese, Bhutanese, Western and Turkic dragons. In Chinese art, dragons are typically portrayed as long, scaled, serpentine creatures with four legs...

 bone
Bone
Bones are rigid organs that constitute part of the endoskeleton of vertebrates. They support, and protect the various organs of the body, produce red and white blood cells and store minerals. Bone tissue is a type of dense connective tissue...

s and documented them as such. For example, Hua Yang Guo Zhi, a book written by Zhang Qu during the Western Jin Dynasty, reported the discovery of dragon bones at Wucheng in Sichuan
Sichuan
' , known formerly in the West by its postal map spellings of Szechwan or Szechuan is a province in Southwest China with its capital in Chengdu...

 Province. Villagers in central China have long unearthed fossilized "dragon bones" for use in traditional medicines, a practice that continues today. In Europe, dinosaur fossils were generally believed to be the remains of giants
Giant (mythology)
The mythology and legends of many different cultures include monsters of human appearance but prodigious size and strength. "Giant" is the English word commonly used for such beings, derived from one of the most famed examples: the gigantes of Greek mythology.In various Indo-European mythologies,...

 and other creatures killed by the Great Flood.

Scholarly descriptions of what would now be recognized as dinosaur bones first appeared in the late 17th century in England
England
England is a country that is part of the United Kingdom. It shares land borders with Scotland to the north and Wales to the west; the Irish Sea is to the north west, the Celtic Sea to the south west, with the North Sea to the east and the English Channel to the south separating it from continental...

. Part of a bone, now known to have been the femur
Femur
The femur , or thigh bone, is the most proximal bone of the leg in tetrapod vertebrates capable of walking or jumping, such as most land mammals, birds, many reptiles such as lizards, and amphibians such as frogs. In vertebrates with four legs such as dogs and horses, the femur is found only in...

 of a Megalosaurus
Megalosaurus
Megalosaurus is a genus of large meat-eating theropod dinosaurs of the Middle Jurassic period of Europe...

, was recovered from a limestone quarry at Cornwell near Chipping Norton, Oxfordshire
Oxfordshire
Oxfordshire is a county in the South East region of England, bordering on Warwickshire and Northamptonshire , Buckinghamshire , Berkshire , Wiltshire and Gloucestershire ....

, England, in 1676. The fragment was sent to Robert Plot
Robert Plot
Robert Plot was an English naturalist, first Professor of Chemistry at the University of Oxford, and the first keeper of the Ashmolean Museum....

, Professor of Chemistry at the University of Oxford
University of Oxford
The University of Oxford is a university located in Oxford, United Kingdom. It is the second-oldest surviving university in the world and the oldest in the English-speaking world. Although its exact date of foundation is unclear, there is evidence of teaching as far back as 1096...

 and first curator of the Ashmolean Museum
Ashmolean Museum
The Ashmolean Museum on Beaumont Street, Oxford, England, is the world's first university museum...

, who published a description in his Natural History of Oxfordshire in 1677. He correctly identified the bone as the lower extremity of the femur
Femur
The femur , or thigh bone, is the most proximal bone of the leg in tetrapod vertebrates capable of walking or jumping, such as most land mammals, birds, many reptiles such as lizards, and amphibians such as frogs. In vertebrates with four legs such as dogs and horses, the femur is found only in...

 of a large animal, and recognized that it was too large to belong to any known species. He therefore concluded it to be the thigh bone of a giant human similar to those mentioned in the Bible
Bible
The Bible refers to any one of the collections of the primary religious texts of Judaism and Christianity. There is no common version of the Bible, as the individual books , their contents and their order vary among denominations...

. In 1699, Edward Lhuyd
Edward Lhuyd
Edward Lhuyd was a Welsh naturalist, botanist, linguist, geographer and antiquary. He is also known by the Latinized form of his name, Eduardus Luidius....

, a friend of Sir Isaac Newton, was responsible for the first published scientific treatment of what would now be recognized as a dinosaur when he described and named a sauropod tooth
Tooth
Teeth are small, calcified, whitish structures found in the jaws of many vertebrates that are used to break down food. Some animals, particularly carnivores, also use teeth for hunting or for defensive purposes. The roots of teeth are embedded in the Mandible bone or the Maxillary bone and are...

, "Rutellum implicatum
Rutellum
Rutellum is the pre-Linnaean name given to a dinosaur specimen from the Middle Jurassic. It was a sauropod, possibly a cetiosaurid, which lived in what is now England. The specimen, called Rutellum implicatum, was described in 1699 by Edward Lhuyd, and is notable as the earliest named entity that...

", that had been found in Caswell, near Witney, Oxfordshire.
Between 1815 and 1824, the Rev William Buckland
William Buckland
The Very Rev. Dr William Buckland DD FRS was an English geologist, palaeontologist and Dean of Westminster, who wrote the first full account of a fossil dinosaur, which he named Megalosaurus...

, a professor of geology
Geology
Geology is the science comprising the study of solid Earth, the rocks of which it is composed, and the processes by which it evolves. Geology gives insight into the history of the Earth, as it provides the primary evidence for plate tectonics, the evolutionary history of life, and past climates...

 at Oxford University
University of Oxford
The University of Oxford is a university located in Oxford, United Kingdom. It is the second-oldest surviving university in the world and the oldest in the English-speaking world. Although its exact date of foundation is unclear, there is evidence of teaching as far back as 1096...

, collected more fossilized bones of Megalosaurus and became the first person to describe a dinosaur in a scientific journal
Scientific journal
In academic publishing, a scientific journal is a periodical publication intended to further the progress of science, usually by reporting new research. There are thousands of scientific journals in publication, and many more have been published at various points in the past...

. The second dinosaur genus to be identified, Iguanodon
Iguanodon
Iguanodon is a genus of ornithopod dinosaur that lived roughly halfway between the first of the swift bipedal hypsilophodontids and the ornithopods' culmination in the duck-billed dinosaurs...

, was discovered in 1822 by Mary Ann Mantell – the wife of English geologist Gideon Mantell
Gideon Mantell
Gideon Algernon Mantell MRCS FRS was an English obstetrician, geologist and palaeontologist...

. Gideon Mantell recognized similarities between his fossils and the bones of modern iguana
Iguana
Iguana is a herbivorous genus of lizard native to tropical areas of Central America and the Caribbean. The genus was first described in 1768 by Austrian naturalist Josephus Nicolaus Laurenti in his book Specimen Medicum, Exhibens Synopsin Reptilium Emendatam cum Experimentis circa Venena...

s. 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 1842 the English paleontologist Richard Owen
Richard Owen
Sir Richard Owen, FRS KCB was an English biologist, comparative anatomist and palaeontologist.Owen is probably best remembered today for coining the word Dinosauria and for his outspoken opposition to Charles Darwin's theory of evolution by natural selection...

 coined the term "dinosaur". He recognized that the remains that had been found so far, Iguanodon, Megalosaurus and Hylaeosaurus
Hylaeosaurus
Hylaeosaurus is the most obscure of the three animals used by Sir Richard Owen to first define the new group Dinosauria, in 1842. The original specimen, recovered by Gideon Mantell from the Tilgate Forest in the south of England in 1832, now resides in the Natural History Museum of London, where...

, shared a number of distinctive features, and so decided to present them as a distinct taxonomic group. With the backing of Prince Albert of Saxe-Coburg-Gotha, the husband of Queen Victoria
Victoria of the United Kingdom
Victoria was the monarch of the United Kingdom of Great Britain and Ireland from 20 June 1837 until her death. From 1 May 1876, she used the additional title of Empress of India....

, Owen established the Natural History Museum
Natural History Museum
The Natural History Museum is one of three large museums on Exhibition Road, South Kensington, London, England . Its main frontage is on Cromwell Road...

 in South Kensington
South Kensington
South Kensington is a district in the Royal Borough of Kensington and Chelsea in London. It is a built-up area located 2.4 miles west south-west of Charing Cross....

, London
London
London is the capital city of :England and the :United Kingdom, the largest metropolitan area in the United Kingdom, and the largest urban zone in the European Union by most measures. Located on the River Thames, London has been a major settlement for two millennia, its history going back to its...

, to display the national collection of dinosaur fossils and other biological and geological exhibits.

In 1858, the first known American dinosaur was discovered, in marl
Marl
Marl or marlstone is a calcium carbonate or lime-rich mud or mudstone which contains variable amounts of clays and aragonite. Marl was originally an old term loosely applied to a variety of materials, most of which occur as loose, earthy deposits consisting chiefly of an intimate mixture of clay...

 pits in the small town of Haddonfield, New Jersey
Haddonfield, New Jersey
Haddonfield is a borough located in Camden County, New Jersey. As of the 2010 United States Census, the borough had a total population of 11,593....

 (although fossils had been found before, their nature had not been correctly discerned). The creature was named Hadrosaurus
Hadrosaurus
Hadrosaurus is a valid genus of hadrosaurid dinosaur. In 1858, a skeleton of a dinosaur from this genus was the first dinosaur skeleton known from more than isolated teeth to be found in North America. In 1868, it became the first ever mounted dinosaur skeleton...

 foulkii
. It was an extremely important find: Hadrosaurus was one of the first nearly complete dinosaur skeletons found (the first was in 1834, in Maidstone, Kent, England), and it was clearly a bipedal creature. This was a revolutionary discovery as, until that point, most scientists had believed dinosaurs walked on four feet, like other lizards. Foulke's discoveries sparked a wave of dinosaur mania in the United States
United States
The United States of America is a federal constitutional republic comprising fifty states and a federal district...

.
Dinosaur mania was exemplified by the fierce rivalry between Edward Drinker Cope
Edward Drinker Cope
Edward Drinker Cope was an American paleontologist and comparative anatomist, as well as a noted herpetologist and ichthyologist. Born to a wealthy Quaker family, Cope distinguished himself as a child prodigy interested in science; he published his first scientific paper at the age of nineteen...

 and Othniel Charles Marsh
Othniel Charles Marsh
Othniel Charles Marsh was an American paleontologist. Marsh was one of the preeminent scientists in the field; the discovery or description of dozens of news species and theories on the origins of birds are among his legacies.Born into a modest family, Marsh was able to afford higher education...

, both of whom raced to be the first to find new dinosaurs in what came to be known as the Bone Wars
Bone Wars
The Bone Wars, also known as the "Great Dinosaur Rush", refers to a period of intense fossil speculation and discovery during the Gilded Age of American history, marked by a heated rivalry between Edward Drinker Cope and Othniel Charles Marsh...

. The feud probably originated when Marsh publicly pointed out that Cope's reconstruction of an Elasmosaurus
Elasmosaurus
Elasmosaurus + σαυρος sauros 'lizard') is a genus of plesiosaur with an extremely long neck that lived in the Late Cretaceous period , 80.5 million years ago.-Description:...

skeleton was flawed: Cope had inadvertently placed the plesiosaur
Plesiosaur
Plesiosauroidea is an extinct clade of carnivorous plesiosaur marine reptiles. Plesiosauroids, are known from the Jurassic and Cretaceous Periods...

's head at what should have been the animal's tail end. The fight between the two scientists lasted for over 30 years, ending in 1897 when Cope died after spending his entire fortune on the dinosaur hunt. Marsh 'won' the contest primarily because he was better funded through a relationship with the US Geological Survey. Unfortunately, many valuable dinosaur specimens were damaged or destroyed due to the pair's rough methods: for example, their diggers often used dynamite
Dynamite
Dynamite is an explosive material based on nitroglycerin, initially using diatomaceous earth , or another absorbent substance such as powdered shells, clay, sawdust, or wood pulp. Dynamites using organic materials such as sawdust are less stable and such use has been generally discontinued...

 to unearth bones (a method modern paleontologists would find appalling). Despite their unrefined methods, the contributions of Cope and Marsh to paleontology were vast: Marsh unearthed 86 new species of dinosaur and Cope discovered 56, a total of 142 new species. Cope's collection is now at the American Museum of Natural History
American Museum of Natural History
The American Museum of Natural History , located on the Upper West Side of Manhattan in New York City, United States, is one of the largest and most celebrated museums in the world...

 in New York
New York
New York is a state in the Northeastern region of the United States. It is the nation's third most populous state. New York is bordered by New Jersey and Pennsylvania to the south, and by Connecticut, Massachusetts and Vermont to the east...

, while Marsh's is on display at the Peabody Museum of Natural History at Yale University
Yale University
Yale University is a private, Ivy League university located in New Haven, Connecticut, United States. Founded in 1701 in the Colony of Connecticut, the university is the third-oldest institution of higher education in the United States...

.

After 1897, the search for dinosaur fossils extended to every continent, including Antarctica. The first Antarctic dinosaur to be discovered, the ankylosaurid Antarctopelta oliveroi
Antarctopelta
Antarctopelta was a genus of ankylosaurian dinosaur with one known species, A. oliveroi, which lived in Antarctica during the Late Cretaceous Period. It was a medium-sized ankylosaur, reaching no more than 4 meters in length, and showed characteristics of two different families, making more...

, was found on Ross Island
Ross Island
Ross Island is an island formed by four volcanoes in the Ross Sea near the continent of Antarctica, off the coast of Victoria Land in McMurdo Sound.-Geography:...

 in 1986, although it was 1994 before an Antarctic species, the theropod Cryolophosaurus ellioti, was formally named and described in a scientific journal.

Current dinosaur "hot spots" include southern South America (especially Argentina
Argentina
Argentina , officially the Argentine Republic , is the second largest country in South America by land area, after Brazil. It is constituted as a federation of 23 provinces and an autonomous city, Buenos Aires...

) and China
China
Chinese civilization may refer to:* China for more general discussion of the country.* Chinese culture* Greater China, the transnational community of ethnic Chinese.* History of China* Sinosphere, the area historically affected by Chinese culture...

. China in particular has produced many exceptional feathered dinosaur specimens due to the unique geology of its dinosaur beds, as well as an ancient arid climate particularly conducive to fossilization.

The "dinosaur renaissance"



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
John Ostrom
John H. Ostrom was an American paleontologist who revolutionized modern understanding of dinosaurs in the 1960s, when he demonstrated that dinosaurs are more like big non-flying birds than they are like lizards , an idea first proposed by Thomas Henry Huxley in the 1860s, but which had garnered...

's discovery of Deinonychus
Deinonychus
Deinonychus was a genus of carnivorous dromaeosaurid dinosaur. There is one described species, Deinonychus antirrhopus. This 3.4 meter long dinosaur lived during the early Cretaceous Period, about 115–108 million years ago . Fossils have been recovered from the U.S...

, an active predator that may have been warm-blooded
Warm-blooded
The term warm-blooded is a colloquial term to describe animal species which have a relatively higher blood temperature, and maintain thermal homeostasis primarily through internal metabolic processes...

, in marked contrast to the then-prevailing image of dinosaurs as sluggish and cold-blooded
Poikilotherm
A poikilotherm is an organism whose internal temperature varies considerably. It is the opposite of a homeotherm, an organism which maintains thermal homeostasis. Usually the variation is a consequence of variation in the ambient environmental temperature...

. Vertebrate paleontology
Vertebrate paleontology
Vertebrate paleontology is a large subfield to paleontology seeking to discover the behavior, reproduction and appearance of extinct animals with vertebrae or a notochord, through the study of their fossilized remains...

 has become a global science
Science
Science is a systematic enterprise that builds and organizes knowledge in the form of testable explanations and predictions about the universe...

. Major new dinosaur discoveries have been made by paleontologists working in previously unexploited regions, including India
India
India , officially the Republic of India , is a country in South Asia. It is the seventh-largest country by geographical area, the second-most populous country with over 1.2 billion people, and the most populous democracy in the world...

, South America, Madagascar
Madagascar
The Republic of Madagascar is an island country located in the Indian Ocean off the southeastern coast of Africa...

, Antarctica, and most significantly China
China
Chinese civilization may refer to:* China for more general discussion of the country.* Chinese culture* Greater China, the transnational community of ethnic Chinese.* History of China* Sinosphere, the area historically affected by Chinese culture...

 (the amazingly well-preserved feathered dinosaurs
Feathered dinosaurs
The realization that dinosaurs are closely related to birds raised the obvious possibility of feathered dinosaurs. Fossils of Archaeopteryx include well-preserved feathers, but it was not until the early 1990s that clearly non-avialan dinosaur fossils were discovered with preserved feathers...

 in China
China
Chinese civilization may refer to:* China for more general discussion of the country.* Chinese culture* Greater China, the transnational community of ethnic Chinese.* History of China* Sinosphere, the area historically affected by Chinese culture...

 have further consolidated the link between dinosaurs and their conjectured living descendants, modern birds). The widespread application of cladistics
Cladistics
Cladistics is a method of classifying species of organisms into groups called clades, which consist of an ancestor organism and all its descendants . For example, birds, dinosaurs, crocodiles, and all descendants of their most recent common ancestor form a clade...

, 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.

Cultural depictions



By human standards, dinosaurs were creatures of fantastic appearance and often enormous size. As such, they have captured the popular imagination and become an enduring part of human culture. Entry of the word "dinosaur" into the common vernacular
Vernacular
A vernacular is the native language or native dialect of a specific population, as opposed to a language of wider communication that is not native to the population, such as a national language or lingua franca.- Etymology :The term is not a recent one...

 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
Victorian era
The Victorian era of British history was the period of Queen Victoria's reign from 20 June 1837 until her death on 22 January 1901. It was a long period of peace, prosperity, refined sensibilities and national self-confidence...

 England, where in 1854, three decades after the first scientific descriptions of dinosaur remains, the famous dinosaur sculptures
Crystal Palace Dinosaurs
The Crystal Palace Dinosaurs, also known as Dinosaur Court, are a series of sculptures of dinosaurs and extinct mammals located in Crystal Palace, London. Commissioned in 1852 and unveiled in 1854, they were the first dinosaur sculptures in the world, pre-dating the publication of Charles Darwin's...

 were unveiled in London
London
London is the capital city of :England and the :United Kingdom, the largest metropolitan area in the United Kingdom, and the largest urban zone in the European Union by most measures. Located on the River Thames, London has been a major settlement for two millennia, its history going back to its...

's Crystal Palace Park. The Crystal Palace dinosaurs proved so popular that a strong market in smaller replicas soon developed. In subsequent decades, dinosaur exhibits opened at parks and museums around the world, ensuring that successive generations would be introduced to the animals in an immersive and exciting way. Dinosaurs' enduring popularity, in its turn, has resulted in significant public funding for dinosaur science, and has frequently spurred new discoveries. In the United States, for example, the competition between museums for public attention led directly to the Bone Wars
Bone Wars
The Bone Wars, also known as the "Great Dinosaur Rush", refers to a period of intense fossil speculation and discovery during the Gilded Age of American history, marked by a heated rivalry between Edward Drinker Cope and Othniel Charles Marsh...

 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
Literature
Literature is the art of written works, and is not bound to published sources...

, film
Film
A film, also called a movie or motion picture, is a series of still or moving images. It is produced by recording photographic images with cameras, or by creating images using animation techniques or visual effects...

 and other media
Media (communication)
In communications, media are the storage and transmission channels or tools used to store and deliver information or data...

. Beginning in 1852 with a passing mention in Charles Dickens'
Charles Dickens
Charles John Huffam Dickens was an English novelist, generally considered the greatest of the Victorian period. Dickens enjoyed a wider popularity and fame than had any previous author during his lifetime, and he remains popular, having been responsible for some of English literature's most iconic...

 Bleak House
Bleak House
Bleak House is the ninth novel by Charles Dickens, published in twenty monthly installments between March 1852 and September 1853. It is held to be one of Dickens's finest novels, containing one of the most vast, complex and engaging arrays of minor characters and sub-plots in his entire canon...

, dinosaurs have been featured in large numbers of fiction
Fiction
Fiction is the form of any narrative or informative work that deals, in part or in whole, with information or events that are not factual, but rather, imaginary—that is, invented by the author. Although fiction describes a major branch of literary work, it may also refer to theatrical,...

al works. Sir Arthur Conan Doyle's 1912 book The Lost World, the iconic 1933 film King Kong
King Kong (1933 film)
King Kong is a Pre-Code 1933 fantasy monster adventure film co-directed by Merian C. Cooper and Ernest B. Schoedsack, and written by Ruth Rose and James Ashmore Creelman after a story by Cooper and Edgar Wallace. The film tells of a gigantic island-dwelling apeman creature called Kong who dies in...

, 1954's Godzilla
Godzilla (1954 film)
is a 1954 Japanese science fiction film directed by Ishirō Honda and produced by Tomoyuki Tanaka. The film stars Akira Takarada, Momoko Kōchi, Akihiko Hirata and Takashi Shimura. The film tells the story of Godzilla, a giant monster mutated by nuclear radiation, who ravages Japan, bringing back the...

and its many sequels, the best-selling 1990 novel Jurassic Park by Michael Crichton
Michael Crichton
John Michael Crichton , best known as Michael Crichton, was an American best-selling author, producer, director, and screenwriter, best known for his work in the science fiction, medical fiction, and thriller genres. His books have sold over 200 million copies worldwide, and many have been adapted...

 and its 1993 film adaptation
Jurassic Park (film)
Jurassic Park is a 1993 American science fiction adventure film directed by Steven Spielberg. The film is based on the novel of the same name by Michael Crichton. It stars Sam Neill, Laura Dern, Jeff Goldblum, Richard Attenborough, Martin Ferrero, and Bob Peck...

 are just a few notable examples of dinosaur appearances in fiction. Authors of general-interest non-fiction
Non-fiction
Non-fiction is the form of any narrative, account, or other communicative work whose assertions and descriptions are understood to be fact...

 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
Advertising
Advertising is a form of communication used to persuade an audience to take some action with respect to products, ideas, or services. Most commonly, the desired result is to drive consumer behavior with respect to a commercial offering, although political and ideological advertising is also common...

; numerous companies have referenced dinosaurs in printed or televised advertisements, either in order to sell their own products or in order to characterize their rivals as slow-moving, dim-witted or obsolete.

See also


Further reading

  • Bakker, Robert T. (1986). The Dinosaur Heresies: New Theories Unlocking the Mystery of the Dinosaurs and Their Extinction
    The Dinosaur Heresies
    The Dinosaur Heresies: New Theories Unlocking the Mystery of the Dinosaurs and Their Extinction was a 1986 book published by Robert T. Bakker, a prominent paleontologist....

    . New York: Morrow. ISBN 0-688-04287-2.
  • Holtz, Thomas R. Jr. (2007). Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages
    Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages
    Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages is a book by Dr. Thomas R. Holtz, Jr. & Luis Rey. It was published in 2007 by Random House.-Table of contents:* Introduction. The world of dinosaurs...

    . New York: Random House. ISBN 978-0-375-82419-7.
  • Paul, Gregory S.
    Gregory S. Paul
    Gregory Scott Paul is a freelance researcher, author and illustrator who works in paleontology, and more recently has examined sociology and theology. He is best known for his work and research on theropod dinosaurs and his detailed illustrations, both live and skeletal...

     (2000). The Scientific American Book of Dinosaurs. New York: St. Martin's Press. ISBN 0-312-26226-4.
  • Paul, Gregory S. (2002). Dinosaurs of the Air: The Evolution and Loss of Flight in Dinosaurs and Birds. Baltimore: The Johns Hopkins University Press. ISBN 0-8018-6763-0.

External links



General

Images


Video

Popular
  • Dinosaurnews (www.dinosaurnews.org) The dinosaur-related headlines from around the world. Recent news on dinosaurs, including finds and discoveries, and many links.
  • Dinosauria From UC Berkeley Museum of Paleontology Detailed information – scroll down for menu.
  • LiveScience.com All about dinosaurs, with current featured articles.
  • Zoom Dinosaurs (www.enchantedlearning.com) From Enchanted Learning. Kids' site, info pages and stats, theories, history.
  • Dinosaur genus list contains data tables on nearly every published Mesozoic dinosaur genus as of January 2011.
  • LiveScience.com Giant Dinosaurs Get Downsized by LiveScience, June 21, 2009


Technical
  • Palaeontologia Electronica From Coquina Press. Online technical journal.
  • Dinobase A searchable dinosaur database, from the University of Bristol, with dinosaur lists, classification, pictures, and more.
  • DinoData (www.dinodata.org) Technical site, essays, classification, anatomy.
  • Dinosauria On-Line (www.dinosauria.com) Technical site, essays, pronunciation, dictionary.
  • Thescelosaurus! By Justin Tweet. Includes a cladogram and small essays on each relevant genera and species.
  • Dinosauromorpha Cladogram From Palaeos. A detailed amateur site about all things paleo.