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Dinosaurs are a diverse group of animals of the clade Dinosauria. They first appeared during the Triassic period, 231.4 million years ago, and were the dominant terrestrial vertebrates for 135 million years, from the beginning of the Jurassic (about 201 million years ago) until the end of the Cretaceous (66 million years ago), when the Cretaceous–Paleogene extinction event led to the extinction of most dinosaur groups at the close of the Mesozoic Era. The fossil record indicates that birds evolved from theropod dinosaurs during the Jurassic Period and, consequently, they are considered a subgroup of dinosaurs by many paleontologists.[1] Some birds survived the extinction event that occurred 66 million years ago, and their descendants continue the dinosaur lineage to the present day.[2]

Dinosaurs are a varied group of animals from taxonomic, morphological and ecological standpoints. Birds, at over 10,000 living species,[3] are the most diverse group of vertebrates besides perciform fish.[4] Using fossil evidence, paleontologists have identified over 500 distinct genera[5] and more than 1,000 different species of non-avian dinosaurs.[6] Dinosaurs are represented on every continent by both extant species and fossil remains.[7] Some are herbivorous, others carnivorous. While dinosaurs were ancestrally bipedal, many extinct groups included quadrupedal species, and some were able to shift between these stances. Elaborate display structures such as horns or crests are common to all dinosaur groups, and some extinct groups developed skeletal modifications such as bony armor and spines. Evidence suggests that egg laying and nest building are additional traits shared by all dinosaurs. While modern dinosaurs (birds) are generally small due to the constraints of flight, many prehistoric dinosaurs were large-bodied—the largest sauropod dinosaurs may have achieved lengths of 58 meters (190 feet) and heights of 9.25 meters (30 feet 4 inches).[8] Still, the idea that non-avian dinosaurs were uniformly gigantic is a misconception based on preservation bias, as large, sturdy bones are more likely to last until they are fossilized. Many dinosaurs were quite small: Xixianykus, for example, was only about 50 cm (20 in) long.

Although the word dinosaur means "terrible lizard", the name is somewhat misleading, as dinosaurs are not lizards. Instead, they represent a separate group of reptiles that, like many extinct forms, did not exhibit characteristics traditionally seen as reptilian, such as a sprawling limb posture or ectothermy. Additionally, many prehistoric animals, including mosasaurs, ichthyosaurs, pterosaurs, plesiosaurs, and Dimetrodon, are popularly conceived of as dinosaurs, but are not taxonomically classified as dinosaurs. Through the first half of the 20th century, before birds were recognized to be dinosaurs, most of the scientific community believed dinosaurs to have been sluggish and cold-blooded. Most research conducted since the 1970s, however, has indicated that all dinosaurs were active animals with elevated metabolisms and numerous adaptations for social interaction.

Since the first dinosaur fossils were recognized in the early 19th century, mounted fossil dinosaur skeletons have been major attractions at museums around the world, and dinosaurs have become an enduring part of world culture. The large sizes of some groups, as well as their seemingly monstrous and fantastic nature, have ensured dinosaurs' regular appearance in best-selling books and films, such as Jurassic Park. Persistent public enthusiasm for the animals has resulted in significant funding for dinosaur science, and new discoveries are regularly covered by the media.

Contents [hide] 1 Etymology 2 Definition 2.1 General description 2.2 Distinguishing anatomical features 3 Evolutionary history 3.1 Origins and early evolution 3.2 Evolution and paleobiogeography 4 Classification 4.1 Taxonomy 5 Biology 5.1 Size 5.1.1 Largest and smallest 5.2 Behavior 5.3 Communication 5.4 Reproductive biology 5.5 Physiology 6 Origin of birds 6.1 Feathers 6.2 Skeleton 6.3 Soft anatomy 6.4 Behavioral evidence 7 Extinction of major groups 7.1 Impact event 7.2 Deccan Traps 7.3 Possible Paleocene survivors 8 History of study 8.1 "Dinosaur renaissance" 8.2 Soft tissue and DNA 8.3 Timeline of notable dinosaur taxonomic descriptions 9 Cultural depictions 10 See also 11 Notes and references 12 Further reading 13 External links Etymology The taxon Dinosauria was formally named in 1842 by paleontologist Sir Richard Owen, who used it to refer to the "distinct tribe or sub-order of Saurian Reptiles" that were then being recognized in England and around the world.[9] The term is derived from the Greek words δεινός (deinos, meaning "terrible," "potent," or "fearfully great") and σαῦρος (sauros, meaning "lizard" or "reptile").[9][10] 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.[11]

Definition

Triceratops horridus skeleton, Natural History Museum of Los Angeles County Under phylogenetic taxonomy, dinosaurs are usually defined as the group consisting of Triceratops, Neornithes [modern birds], their most recent common ancestor (MRCA), and all descendants.[12] It has also been suggested that Dinosauria be defined with respect to the MRCA of Megalosaurus and Iguanodon, because these were two of the three genera cited by Richard Owen when he recognized the Dinosauria.[13] Both definitions result in the same set of animals being defined as dinosaurs: "Dinosauria = Ornithischia + Saurischia", encompassing theropods (mostly bipedal carnivores and birds), ankylosaurians (armored herbivorous quadrupeds), stegosaurians (plated herbivorous quadrupeds), ceratopsians (herbivorous quadrupeds with horns and frills), ornithopods (bipedal or quadrupedal herbivores including "duck-bills"), and sauropodomorphs (mostly large herbivorous quadrupeds with long necks and tails).[14]

small sparrow with pale belly and breast and patterned wing and head The common House sparrow (Passer domesticus) is often used to represent modern birds in definitions of the group Dinosauria. Birds are now recognized as being the sole surviving lineage of theropod dinosaurs. In traditional taxonomy, birds were considered a separate class that had evolved from dinosaurs, a distinct superorder. However, a majority of contemporary paleontologists concerned with dinosaurs reject the traditional style of classification in favor of phylogenetic nomenclature; this approach requires that, for a group to be natural, all descendants of members of the group must be included in the group as well. Birds are thus considered to be dinosaurs and dinosaurs are, therefore, not extinct. Birds are classified as belonging to the subgroup Maniraptora, which are coelurosaurs, which are theropods, which are saurischians, which are dinosaurs.[15]

General description Using one of the above definitions, dinosaurs can be generally described as archosaurs with hind limbs held erect beneath the body.[16] Many prehistoric animal groups are popularly conceived of as dinosaurs, such as ichthyosaurs, mosasaurs, plesiosaurs, pterosaurs, and Dimetrodon, but are not classified scientifically as dinosaurs, and none had the erect hind limb posture characteristic of true dinosaurs.[17] Dinosaurs were the dominant terrestrial vertebrates of the Mesozoic, especially the Jurassic and Cretaceous periods. Other groups of animals were restricted in size and niches; mammals, for example, rarely exceeded the size of a cat, and were generally rodent-sized carnivores of small prey.[18]

Dinosaurs have always been an extremely varied group of animals; according to a 2006 study, over 500 non-avialan 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.[5] An earlier study predicted that about 3400 dinosaur genera existed, including many which would not have been preserved in the fossil record.[19] By September 17, 2008, 1047 different species of dinosaurs had been named.[6] Some are herbivorous, others carnivorous, including seed-eaters, fish-eaters, insectivores, and omnivores. While dinosaurs were ancestrally bipedal (as are all modern birds), some prehistoric species were quadrupeds, and others, such as Ammosaurus and Iguanodon, could walk just as easily on two or four legs. Cranial modifications like horns and crests are common dinosaurian traits, and some extinct species had bony armor. Although known for large size, many Mesozoic dinosaurs were human-sized or smaller, and modern birds are generally small in size. Dinosaurs today inhabit every continent, and fossils show that they had achieved global distribution by at least the early Jurassic period.[7] Modern birds inhabit most available habitats, from terrestrial to marine, and there is evidence that some non-avialan dinosaurs (such as Microraptor) could fly or at least glide, and others, such as spinosaurids, had semi-aquatic habits.[20]

Distinguishing anatomical features While recent discoveries have made it more difficult to present a universally agreed-upon list of dinosaurs' distinguishing features, nearly all dinosaurs discovered so far share certain modifications to the ancestral archosaurian skeleton, or are clear descendants of older dinosaurs showing these modifications. Although some later groups of dinosaurs featured further modified versions of these traits, they are considered typical for Dinosauria; the earliest dinosaurs had them and passed them on to their descendants. Such modifications, originating in the last common ancestor of a certain taxonomic group, are called the synapomorphies of such a group.[21]

A detailed assessment of archosaur interrelations by S. Nesbitt[22] confirmed or found the following twelve unambiguous synapomorphies, some previously known:

in the skull, a supratemporal fossa (excavation) is present in front of the supratemporal fenestra, the main opening in the rear skull roof epipophyses, obliquely backward pointing processes on the rear top corners, present in the anterior (front) neck vertebrae behind the atlas and axis, the first two neck vertebrae apex of deltopectoral crest (a projection on which the deltopectoral muscles attach) located at or more than 30% down the length of the humerus (upper arm bone) radius, a lower arm bone, shorter than 80% of humerus length fourth trochanter (projection where the caudofemoralis muscle attaches on the inner rear shaft) on the femur (thighbone) is a sharp flange fourth trochanter asymmetrical, with distal, lower, margin forming a steeper angle to the shaft on the astragalus and calcaneum, upper ankle bones, the proximal articular facet, the top connecting surface, for the fibula occupies less than 30% of the transverse width of the element exoccipitals (bones at the back of the skull) do not meet along the midline on the floor of the endocranial cavity, the inner space of the braincase in the pelvis, the proximal articular surfaces of the ischium with the ilium and the pubis are separated by a large concave surface (on the upper side of the ischium a part of the open hip joint is located between the contacts with the pubic bone and the ilium) cnemial crest on the tibia (protruding part of the top surface of the shinbone) arcs anterolaterally (curves to the front and the outer side) distinct proximodistally oriented (vertical) ridge present on the posterior face of the distal end of the tibia (the rear surface of the lower end of the shinbone) Concave articular surface for the fibula of the calcaneum (the top surface of the calcaneum, where it touches the fibula, has a hollow profile) Nesbitt found a number of further potential synapomorphies, and discounted a number of synapomorphies previously suggested. Some of these are also present in silesaurids, which Nesbitt recovered as a sister group to Dinosauria, including a large anterior trochanter, metatarsals II and IV of subequal length, reduced contact between ischium and pubis, the presence of a cnemial crest on the tibia and of an ascending process on the astragalus, and many others.[12]

Diagram of a typical diapsid skull

Hip joints and hindlimb postures of: (left to right) typical reptiles (sprawling), dinosaurs and mammals (erect), and rauisuchians (erect) A variety of other skeletal features are shared by dinosaurs. However, because they are either common to other groups of archosaurs or were not present in all early dinosaurs, these features are not considered to be synapomorphies. For example, as diapsids, dinosaurs ancestrally had two pairs of temporal fenestrae (openings in the skull behind the eyes), and as members of the diapsid group Archosauria, had additional openings in the snout and lower jaw.[23] Additionally, several characteristics once thought to be synapomorphies are now known to have appeared before dinosaurs, or were absent in the earliest dinosaurs and independently evolved by different dinosaur groups. These include an elongated scapula, or shoulder blade; a sacrum composed of three or more fused vertebrae (three are found in some other archosaurs, but only two are found in Herrerasaurus);[12] and a perforate acetabulum, or hip socket, with a hole at the center of its inside surface (closed in Saturnalia, for example).[24][25] 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.[26]

Dinosaurs stand with their hind limbs erect in a manner similar to most modern mammals, but distinct from most other reptiles, whose limbs sprawl out to either side.[27] This posture is due to the development of a laterally facing recess in the pelvis (usually an open socket) and a corresponding inwardly facing distinct head on the femur.[28] Their erect posture enabled early dinosaurs to breathe easily while moving, which likely permitted stamina and activity levels that surpassed those of "sprawling" reptiles.[29] Erect limbs probably also helped support the evolution of large size by reducing bending stresses on limbs.[30] Some non-dinosaurian archosaurs, including rauisuchians, also had erect limbs but achieved this by a "pillar erect" configuration of the hip joint, where instead of having a projection from the femur insert on a socket on the hip, the upper pelvic bone was rotated to form an overhanging shelf.[30]

Evolutionary history Main article: Evolution of dinosaurs Origins and early evolution

Skeleton of Marasuchus lilloensis, a dinosaur-like ornithodiran Full skeleton of an early carnivorous dinosaur, displayed in a glass case in a museum The early forms Herrerasaurus (large), Eoraptor (small) and a Plateosaurus skull Dinosaurs diverged from their archosaur ancestors during the Middle to Late Triassic period, roughly 20 million years after the Permian–Triassic extinction event wiped out an estimated 95% of all life on Earth.[31][32] Radiometric dating of the rock formation that contained fossils from the early dinosaur genus Eoraptor at 231.4 million years old establishes its presence in the fossil record at this time.[33] Paleontologists think that Eoraptor resembles the common ancestor of all dinosaurs;[34] if this is true, its traits suggest that the first dinosaurs were small, bipedal predators.[35] The discovery of primitive, dinosaur-like ornithodirans such as Marasuchus and Lagerpeton in Argentinian Middle Triassic strata supports this view; analysis of recovered fossils suggests that these animals were indeed small, bipedal predators. Dinosaurs may have appeared as early as 243 million years ago, as evidenced by remains of the genus Nyasasaurus from that period, though known fossils of these animals are too fragmentary to tell if they are dinosaurs or very close dinosaurian relatives.[36]

When dinosaurs appeared, they were not the dominant terrestrial animals. The terrestrial habitats were occupied by various types of archosauromorphs and therapsids, like cynodonts and rhynchosaurs. Their main competitors were the pseudosuchia, such as aetosaurs, ornithosuchids and rauisuchians, which were more successful than the dinosaurs.[37] Most of these other animals became extinct in the Triassic, in one of two events. First, at about 215 million years ago, variety of basal archosauromorphs, including the protorosaurs, 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, phytosaurs, and rauisuchians. Rhynchosaurs and dicynodonts survived (at least in some areas) at least as late as early-mid Norian and early Rhaetian, respectively,[38][39] and the exact date of their extinction is uncertain. These losses left behind a land fauna of crocodylomorphs, dinosaurs, mammals, pterosaurians, and turtles.[12] The first few lines of early dinosaurs diversified through the Carnian and Norian stages of the Triassic, possibly by occupying the niches of the groups that became extinct.[14]

Evolution and paleobiogeography Dinosaur evolution after the Triassic follows changes in vegetation and the location of continents. In the Late Triassic and Early Jurassic, the continents were connected as the single landmass Pangaea, and there was a worldwide dinosaur fauna mostly composed of coelophysoid carnivores and early sauropodomorph herbivores.[40] Gymnosperm plants (particularly conifers), a potential food source, radiated in the Late Triassic. Early sauropodomorphs did not have sophisticated mechanisms for processing food in the mouth, and so must have employed other means of breaking down food farther along the digestive tract.[41] The general homogeneity of dinosaurian faunas continued into the Middle and Late Jurassic, where most localities had predators consisting of ceratosaurians, spinosauroids, and carnosaurians, and herbivores consisting of stegosaurian ornithischians and large sauropods. Examples of this include the Morrison Formation of North America and Tendaguru Beds of Tanzania. Dinosaurs in China show some differences, with specialized sinraptorid theropods and unusual, long-necked sauropods like Mamenchisaurus.[40] Ankylosaurians and ornithopods were also becoming more common, but prosauropods had become extinct. Conifers and pteridophytes were the most common plants. Sauropods, like the earlier prosauropods, were not oral processors, but ornithischians were evolving various means of dealing with food in the mouth, including potential cheek-like organs to keep food in the mouth, and jaw motions to grind food.[41] Another notable evolutionary event of the Jurassic was the appearance of true birds, descended from maniraptoran coelurosaurians.[42]

By the Early Cretaceous and the ongoing breakup of Pangaea, dinosaurs were becoming strongly differentiated by landmass. The earliest part of this time saw the spread of ankylosaurians, iguanodontians, and brachiosaurids through Europe, North America, and northern Africa. These were later supplemented or replaced in Africa by large spinosaurid and carcharodontosaurid theropods, and rebbachisaurid and titanosaurian sauropods, also found in South America. In Asia, maniraptoran coelurosaurians like dromaeosaurids, troodontids, and oviraptorosaurians became the common theropods, and ankylosaurids and early ceratopsians like Psittacosaurus became important herbivores. Meanwhile, Australia was home to a fauna of basal ankylosaurians, hypsilophodonts, and iguanodontians.[40] The stegosaurians appear to have gone extinct at some point in the late Early Cretaceous or early Late Cretaceous. A major change in the Early Cretaceous, which would be amplified in the Late Cretaceous, was the evolution of flowering plants. At the same time, several groups of dinosaurian herbivores evolved more sophisticated ways to orally process food. Ceratopsians developed a method of slicing with teeth stacked on each other in batteries, and iguanodontians refined a method of grinding with tooth batteries, taken to its extreme in hadrosaurids.[41] Some sauropods also evolved tooth batteries, best exemplified by the rebbachisaurid Nigersaurus.[43]

There were three general dinosaur faunas in the Late Cretaceous. In the northern continents of North America and Asia, the major theropods were tyrannosaurids and various types of smaller maniraptoran theropods, with a predominantly ornithischian herbivore assemblage of hadrosaurids, ceratopsians, ankylosaurids, and pachycephalosaurians. In the southern continents that had made up the now-splitting Gondwana, abelisaurids were the common theropods, and titanosaurian sauropods the common herbivores. Finally, in Europe, dromaeosaurids, rhabdodontid iguanodontians, nodosaurid ankylosaurians, and titanosaurian sauropods were prevalent.[40] Flowering plants were greatly radiating,[41] with the first grasses appearing by the end of the Cretaceous.[44] Grinding hadrosaurids and shearing ceratopsians became extremely diverse across North America and Asia. Theropods were also radiating as herbivores or omnivores, with therizinosaurians and ornithomimosaurians becoming common.[41]

The Cretaceous–Paleogene extinction event, which occurred approximately 66 million years ago at the end of the Cretaceous period, caused the extinction of all dinosaur groups except for the neornithine birds. Some other diapsid groups, such as crocodilians, sebecosuchians, turtles, lizards, snakes, sphenodontians, and choristoderans, also survived the event.[45]

The surviving lineages of neornithine birds, including the ancestors of modern ratites, ducks and chickens, and a variety of waterbirds, diversified rapidly at the beginning of the Paleogene period, entering ecological niches left vacant by the extinction of Mesozoic dinosaur groups such as the arboreal enantiornithines, aquatic hesperornithines, and even the larger terrestrial theropods (in the form of Gastornis, mihirungs, and "terror birds"). However, mammals were also rapidly diversifying during this time, and out-competed the neornithines for dominance of most terrestrial niches.[46]

Classification Main article: Dinosaur classification Dinosaurs are archosaurs, like modern crocodilians. Within the archosaur group, dinosaurs are differentiated most noticeably by their gait. Dinosaur legs extend directly beneath the body, whereas the legs of lizards and crocodilians sprawl out to either side.[21]

Collectively, dinosaurs as a clade are divided into two primary branches, Saurischia and Ornithischia. Saurischia includes those taxa sharing a more recent common ancestor with birds than with Ornithischia, while Ornithischia includes all taxa sharing a more recent common ancestor with Triceratops than with Saurischia. Anatomically, these two groups can be distinguished most noticeably by their pelvic structure. Early saurischians—"lizard-hipped", from the Greek sauros (σαῦρος) meaning "lizard" and ischion (ἰσχίον) meaning "hip joint—retained the hip structure of their ancestors, with a pubis bone directed cranially, or forward.[28] This basic form was modified by rotating the pubis backward to varying degrees in several groups (Herrerasaurus,[47] therizinosauroids,[48] dromaeosaurids,[49] and birds[42]). Saurischia includes the theropods (exclusively bipedal and with a wide variety of diets) and sauropodomorphs (long-necked herbivores which include advanced, quadrupedal groups).[50][51]

By contrast, ornithischians—"bird-hipped", from the Greek ornitheios (ὀρνίθειος) meaning "of a bird" and ischion (ἰσχίον) meaning "hip joint"—had a pelvis that superficially resembled a bird's pelvis: the pubis bone was oriented caudally (rear-pointing). Unlike birds, the ornithischian pubis also usually had an additional forward-pointing process. Ornithischia includes a variety of species which were primarily herbivores. (NB: the terms "lizard hip" and "bird hip" are misnomers – birds evolved from dinosaurs with "lizard hips".)[21]