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Dinosaurs Fossil range: Triassic – Cretaceous (excluding Aves)
Mounted skeletons of Tyrannosaurus (left) and Apatosaurus (right) at the American Museum of Natural History.
Scientific classification
Kingdom: Animalia Phylum: Chordata Subphylum: Vertebrata Class: Sauropsida Subclass: Diapsida Infraclass: Archosauromorpha Superorder: Dinosauria * Owen, 1842
Orders & Suborders
Ornithischia Cerapoda Thyreophora Saurischia Sauropodomorpha Theropoda

Dinosaurs were vertebrate animals that dominated terrestrial ecosystems for over 160 million years, first appearing approximately 230 million years ago. At the end of the Cretaceous Period, 65 million years ago, a catastrophic extinction event ended the dominance of dinosaurs on land. One group of dinosaurs is known to have survived to the present day: taxonomists believe modern birds are direct descendants of theropod dinosaurs.

Since the first dinosaur fossils were recognized in the early nineteenth century, mounted dinosaur skeletons have become major attractions at museums around the world. Dinosaurs have become a part of world culture and remain consistently popular among children and adults. They have been featured in best-selling books and films, and new discoveries are regularly covered by the media.

The term dinosaur is sometimes used informally to describe other prehistoric reptiles, such as the pelycosaur Dimetrodon, the winged pterosaurs, and the aquatic ichthyosaurs, plesiosaurs and mosasaurs, although none of these were dinosaurs.

Contents 1 What is a dinosaur? 1.1 Definition 1.1.1 Distinguishing features of dinosaurs 1.1.2 Phylogenetic definition 2 Natural history 2.1 Origins and early evolution 2.2 Classification 2.2.1 Order Saurischia 2.2.2 †Order Ornithischia 2.3 Evolution and paleobiogeography 3 Study of dinosaurs 3.1 Size 3.1.1 Largest and smallest dinosaurs 3.2 Behavior 3.3 Areas of controversy 3.3.1 Physiology 3.3.2 Soft tissue and DNA 4 Feathered dinosaurs and the origin of birds 4.1 Feathers 4.2 Skeleton 4.3 Reproductive biology 4.4 Lungs 4.5 Heart and sleeping posture 4.6 Gizzard 5 Extinction 5.1 Impact event 5.2 Deccan Traps 5.3 Possible Paleocene dinosaurs 6 History of discovery 6.1 The "dinosaur renaissance" 7 Dinosaurs in culture 8 Religious views 9 See also 10 Notes and references 11 General references 12 See also 13 External links

What is a dinosaur?

Dinosaurs, excluding birds, can be generally described as terrestrial archosaurs with limbs held erect beneath the body, that existed from the Carnian faunal stage of the Late Triassic to the Maastrichtian stage of the Late Cretaceous.^[1] This excludes many prehistoric animals that are popularly conceived as dinosaurs. Examples include: marine reptiles like ichthyosaurs, mosasaurs, and plesiosaurs, which were neither terrestrial nor archosaurs; pterosaurs, which were not terrestrial; and Dimetrodon, a Permian animal more closely related to mammals.^[2] Dinosaurs were the dominant terrestrial vertebrates of the Mesozoic, especially the Jurassic 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.^[3] One notable exception is Repenomamus giganticus, a 12 kilograms (26 lb) to 14 kilograms (31 lb) triconodont that is known to have eaten small dinosaurs like young Psittacosaurus.^[4]

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 1,850, nearly 75% of which remain to be discovered.^[5] An earlier study predicted that about 3,400 dinosaur genera existed, including many which would not have been preserved in the fossil record.^[6] Some were herbivorous, others carnivorous. Some dinosaurs were bipeds, some were quadrupeds, and others, such as Ammosaurus and Iguanodon, could walk just as easily on two or four legs. Many had bony armor, 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.^[7] Despite their diversity and dominance, however, as noted, dinosaurs (with the exception of birds) did not spread into aquatic or aerial niches.

Definition

The taxon Dinosauria was formally named in 1842 by English palaeontologist 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.^[8] The term is derived from the Greek words δεινός (deinos meaning "terrible", "fearsome", or "formidable") and σαύρα (saura 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.^[9]

Distinguishing features of dinosaurs

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. 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 structures across a taxonomic group are called synapomorphies.

Dinosaur synapomorphies include an elongated crest on the humerus, or upper arm bone, to accommodate the attachment of deltopectoral muscles; a shelf at the rear of the ilium, or main hip bone; a tibia, or shin bone, featuring a broad lower edge and a flange pointing out and to the rear; and an ascending projection on the astragalus, one of the ankle bones, which secures it to the tibia.^[10]

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. Such shared features include a diapsid skull bearing two pairs of holes in the temporal region; holes in the snout and lower jaw (two characteristics shared by other archosaurs); loss of the skull's postfrontal bone; a long neck incorporating an S-shaped curve;^[11] an elongated scapula, or shoulder blade; forelimbs shorter and lighter than hind limbs, coupled to asymmetrical hands; a sacrum composed of three or more fused vertebrae; and an acetabulum, or hip socket, with a hole at the center of its inside surface.^[12]

The open, or "perforate", hip joint described above had significant implications for dinosaur movement and behavior. Most notably, it allowed dinosaur hind limbs to be "underslung", or situated directly beneath the animals' bodies; this, in turn, allowed dinosaurs to stand erect in a manner similar to modern mammals, but distinct from most other reptiles, whose limbs sprawl out to either side.^[13] Vertical limb configuration also enabled dinosaurs to breathe easily while moving, which likely permitted stamina and activity levels that surpassed those of "sprawling" reptiles.

Phylogenetic definition

Under phylogenetic taxonomy, dinosaurs are usually defined as all descendants of the most recent common ancestor of Triceratops and modern birds.^[14] It has also been suggested that Dinosauria be defined as all the descendants of the most recent common ancestor of Megalosaurus and Iguanodon, because these were two of the three genera cited by Richard Owen when he recognized the Dinosauria.^[15] They are divided into Ornithischia (bird-hipped) and Saurischia (lizard-hipped), depending upon pelvic structure. Ornithischian dinosaurs had a four-pronged pelvic configuration, incorporating a caudally-directed (rear-pointing) pubis bone with (most commonly) a forward-pointing process. By contrast, the pelvic structure of saurischian dinosaurs was three-pronged, and featured a pubis bone directed cranially, or forwards, only.^[13] Ornithischia includes all taxa sharing a more recent common ancestor with Triceratops than with Saurischia, while Saurischia includes those taxa sharing a more recent common ancestor with birds than with Ornithischia.

There is an almost universal consensus among paleontologists that birds are the descendants of theropod dinosaurs. Using the strict cladistical definition that all descendants of a single common ancestor are related, modern birds are dinosaurs and dinosaurs are, therefore, not extinct. Modern birds are classified by most paleontologists as belonging to the subgroup Maniraptora, which are coelurosaurs, which are theropods, which are saurischians, which are dinosaurs.^[16]

However, referring to birds as 'avian dinosaurs' and to all other dinosaurs as 'non-avian dinosaurs' is cumbersome. Birds are still referred to as birds, at least in popular usage and among ornithologists. It is also technically correct to refer to birds as a distinct group under the older Linnaean classification system, which accepts paraphyletic taxa that exclude some descendants of a single common ancestor. Paleontologists mostly use cladistics, which classifies birds as dinosaurs, but some biologists of the older generation do not.

For clarity, this article will use 'dinosaur' as a synonym for 'non-avian dinosaur', and 'bird' as a synonym for 'avian dinosaur' (meaning any animal that evolved from the common ancestor of Archaeopteryx and modern birds). The term 'non-avian dinosaur' will be used for emphasis as needed.

Natural history

Origins and early evolution

Dinosaurs diverged from their archosaur ancestors approximately 230 million years ago 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.^{[17] [18]} Radiometric dating of the rock formation that contained fossils from the early dinosaur genus Eoraptor establishes its presence in the fossil record at this time. Paleontologists believe Eoraptor resembles the common ancestor of all dinosaurs;^[19] if this is true, its traits suggest that the first dinosaurs were small, bipedal predators.^[20] 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.

When dinosaurs appeared, terrestrial habitats were occupied by various types of basal archosaurs and therapsids, such as aetosaurs, cynodonts, dicynodonts, ornithosuchids, rauisuchias, and rhynchosaurs. Most of these other animals became extinct in the Triassic, in one of two events. First, at about the boundary between the Carnian and Norian faunal stages (about 215 million years ago), dicynodonts and a variety of basal archosauromorphs, including the prolacertiforms and rynchosaurs, 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. These losses left behind a land fauna of crocodylomorphs, dinosaurs, mammals, pterosaurians, and turtles.^[10]

The first few lines of primitive dinosaurs diversified through the Carnian and Norian stages of the Triassic, most likely by occupying the niches of groups that became extinct. Traditionally, dinosaurs were thought to have replaced the variety of other Triassic land animals by proving superior through competition. This now appears unlikely, for several reasons. Early considerations of dinosaur evolution had dinosaurs as polyphyletic, with multiple groups of unrelated "dinosaurs" evolving due to similar pressures, but dinosaurs are now known to have formed a single group. Early conceptions also had a long, drawn-out period of competition beginning in the Middle Triassic, but more work has shown that dinosaurs did not appear that early and had a sudden diversification. Dinosaurs do not show a pattern of steadily increasing in diversity and numbers, as would be predicted if they were competitively replacing other groups; instead, they were very rare through the Carnian, making up only 1-2% of individuals present in faunas. In the Norian, however, after the extinction of several other groups, they became significant components of faunas, representing 50-90% of individuals. Also, what had been viewed as a key adaptation of dinosaurs, their erect stance, is now known to have present in several contemporaneous groups that were not as successful (aetosaurs, ornithosuchids, rauisuchians, and some groups of crocodylomorphs). Finally, the Late Triassic itself was a time of great upheaval in life, with shifts in plant life, marine life, and climate.^[10]

Classification

Main article: Dinosaur classification

Dinosaurs (including birds) are archosaurs, like modern crocodilians. Archosaurs' diapsid skulls have two holes, called temporal fenestrae, located where the jaw muscles attach. Most reptiles (including birds) are diapsids; mammals, with only one temporal fenestra, are called synapsids; and turtles, with no temporal fenestra, are anapsids. Anatomically, dinosaurs share many other archosaur characteristics, including teeth that grow from sockets rather than as direct extensions of the jawbones. 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 crocodylians sprawl out to either side. All dinosaurs were land animals.

Many other types of reptiles lived at the same time as the dinosaurs. Some of these are commonly, but incorrectly, thought of as dinosaurs, including plesiosaurs (which are not closely related to the dinosaurs) and pterosaurs, which developed separately from reptilian ancestors in the late Triassic period.

Collectively, dinosaurs are usually regarded as a superorder or an unranked clade. They are divided into two orders, the Saurischia and the Ornithischia, on the basis of their hip structure. Saurischians ('lizard-hipped', from the Greek sauros (σαυρος) meaning 'lizard' and ischion (ισχιον) meaning 'hip joint') are dinosaurs that originally retained the hip structure of their ancestors. They include all the theropods (bipedal carnivores) and sauropods (long-necked herbivores). Ornithischians ('bird-hipped', from the Greek ornitheios (ορνιθειος) meaning 'of a bird' and ischion (ισχιον) meaning 'hip joint') is the other dinosaurian order, most of which were quadrupedal herbivores. (NB: the terms "lizard hip" and "bird-hip" are misnomers — birds evolved from dinosaurs with "lizard hips".)

Saurischian pelvis structure (left side)

Tyrannosaurus pelvis (showing saurischian structure - left side)

Ornithischian pelvis structure (left side).

Edmontosaurus pelvis (showing ornithischian structure - left side)

The following is a simplified classification of dinosaur families. A more detailed version can be found at List of dinosaur classifications.

The dagger (†) is used to indicate taxa that are extinct.

Order Saurischia

• †Infraorder Herrerasauria
• Suborder Theropoda
  • †Superfamily Coelophysoidea
  • †Infraorder Ceratosauria
    • Family Ceratosauridae
    • Family Abelisauridae
  • Clade Tetanurae
    • †Superfamily Spinosauroidea
      • Family Megalosauridae
      • Family Spinosauridae
    • †Infraorder Carnosauria
    • Clade Coelurosauria
      • †Superfamily Tyrannosauroidea
      • †Infraorder Ornithomimosauria
      • †Infraorder Segnosauria
      • †Infraorder Oviraptorosauria
      • †Infraorder Deinonychosauria
        • Family Dromaeosauridae
        • Family Troodontidae

• †Suborder Sauropodomorpha
  • Infraorder Prosauropoda
    • Family Riojasauridae
    • Family Plateosauridae
    • Family Massospondylidae
  • Infraorder Sauropoda
    • Family Anchisauridae
    • Family Melanorosauridae
    • Family Blikanasauridae
    • Family Vulcanodontidae
    • Family Cetiosauridae
    • Family Omeisauridae
    • Clade Turiasauria
    • Clade Neosauropoda
      • Superfamily Diplodocoidea
      • Family Camarasauridae
      • Family Brachiosauridae
      • Superfamily Titanosauroidea

†Order Ornithischia

• Family Fabrosauridae
• Suborder Thyreophora
  • Family Scelidosauridae
  • Infraorder Stegosauria
  • Infraorder Ankylosauria
    • Family Nodosauridae
    • Family Ankylosauridae
• Suborder Cerapoda
  • Family Heterodontosauridae
  • Infraorder Ornithopoda
    • Family Hypsilophodontidae
    • Family Iguanodontidae
    • Superfamily Hadrosauroidea
  • Clade Marginocephalia
    • Infraorder Pachycephalosauria
    • Infraorder Ceratopsia
      • Family Psittacosauridae
      • Family Protoceratopsidae
      • Family Ceratopsidae

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, there was a worldwide dinosaur fauna mostly composed of coelophysoid predators and prosauropod herbivores.^[21] Gymnosperm plants (particularly conifers), a potential food source, radiated in the Late Triassic. Prosauropods did not have sophisticated mechanisms for processing food in the mouth, so must have employed other means of breaking down food farther along the digestive tract.^[22] 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.^[21] 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.^[22] Another notable evolutionary event of the Jurassic was the appearance of true birds, descended from maniraptoran coelurosaurians.^[16]

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 ^[21] 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.^[22] Some sauropods also evolved tooth batteries, best exemplified by the rebbachisaurid Nigersaurus.^[23]

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.^[21] Flowering plants were greatly radiating,^[22] with the first grasses appearing by the end of the Cretaceous.^[24] 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.^[22]

The Cretaceous–Tertiary 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 line that had already given rise to the first birds. Some other diapsid groups, such as crocodylians, lizards, snakes, sphenodontians, and choristoderans, also survived the event.^[25]

Study of dinosaurs

Knowledge about dinosaurs is derived from a variety of fossil and non-fossil records, including fossilized bones, feces, trackways, gastroliths, feathers, impressions of skin, internal organs and soft tissues.^[26][27] Many fields of study contribute to our understanding of dinosaurs, including physics, chemistry, biology, and the earth sciences (of which paleontology is a sub-discipline). Two topics of particular interest and study have been dinosaur size and behavior.

Size

Main article: Dinosaur size

While the evidence is incomplete, it is clear that, as a group, dinosaurs were large. Even by dinosaur standards, the sauropods were gigantic. For much of the dinosaur era, the smallest sauropods were larger than anything else in their habitat, and the largest were an order of magnitude more massive than anything else that has since walked the Earth. Giant prehistoric mammals such as the Indricotherium and the Columbian mammoth 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, which reaches up to 173,000 kilograms (381,400 lb) and over 30 meters (98 ft) in length.^[28]

Most dinosaurs, however, were much smaller than the giant sauropods. Current evidence suggests that dinosaur average size varied through the Triassic, early Jurassic, late Jurassic and Cretaceous periods.^[19] Theropod dinosaurs, when sorted by estimated weight into categories based on order of magnitude, most often fall into the 100 kilograms (220 lb) to 1,000 kilograms (2,205 lb) category, whereas Recent predatory carnivorans peak in the 10 kilograms (22 lb) to 100 kilograms (220 lb) category.^[29] A rough estimate for average dinosaur weight is about 100 kilograms (220 lb). This contrasts sharply with the size of Cenozoic mammals, estimated by the same source (the National Museum of Natural History) as about 2 kilograms (4 lb) to 5 kilograms (11 lb).^[30]

Largest and smallest dinosaurs

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. As a result, scientists will probably never be certain of the largest and smallest dinosaurs.

The tallest and heaviest dinosaur known from good skeletons is Brachiosaurus brancai (also known as Giraffatitan). Its remains were discovered in Tanzania between 1907–12. Bones from multiple similarly-sized individuals were incorporated into the skeleton now mounted and on display at the Humboldt Museum of Berlin;^[31] this mount is 12 meters (39 ft) tall and 22.5 meters (74 ft) long, and would have belonged to an animal that weighed between 30,000 kilograms (66,139 lb) to 60,000 kilograms (132,277 lb). The longest complete dinosaur is the 27 m (89 ft) long Diplodocus, which was discovered in Wyoming in the United States and displayed in Pittsburgh's 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 specimens on record were all discovered in the 1970s or later, and include the massive Argentinosaurus, which may have weighed 80,000 kilograms (176,370 lb) to 100,000 kilograms (220,462 lb); the longest, the 40 meters (131 ft)) long Supersaurus; and the tallest, the 18 meters (59 ft) Sauroposeidon, which could have reached a sixth-floor window. The longest of them all may have been Amphicoelias fragillimus, known only from a now lost partial vertebral neural arch described in 1878. Extrapolating from the illustration of this bone, the animal may have been 58 meters (190 ft) long and weighed over 120,000 kilograms (264,555 lb),^[32] heavier than all known dinosaurs except possibly the poorly known Bruhathkayosaurus, which could have weighed 175,000 kilograms (385,809 lb) to 220,000 kilograms (485,017 lb). The largest known carnivorous dinosaur was Spinosaurus, reaching a length of 16 meters (52 ft) to 18 meters (59 ft), and weighing in at 8,150 kilograms (17,968 lb).^[33] Other large meat-eaters included Giganotosaurus, Mapusaurus, Tyrannosaurus rex and Carcharodontosaurus.

Not including modern birds, the smallest dinosaurs known were about the size of a crow or a chicken. The theropods Microraptor and Parvicursor were both under 0.6 meters (2 ft) in length.

Behavior

Interpretations of dinosaur behavior are generally based on the pose of body fossils and their habitat, computer simulations of their biomechanics, and comparisons with modern animals in similar ecological niches. 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 direct evidence of herding behavior was the 1878 discovery of 31 Iguanodon dinosaurs which were thought to have perished together in Bernissart, Belgium, after they fell into a deep, flooded sinkhole and drowned.^[34] Other mass death sites have been subsequently discovered. Those, along with multiple trackways, suggest that herd or pack behavior was common in many dinosaur species. Trackways of hundreds or even thousands of herbivores indicate that duck-bills (hadrosaurids) may have moved in great herds, like the American Bison or the African Springbok. Sauropod tracks document that these animals traveled in groups composed of several different species, at least in Oxford, England,^[35] and others kept their young in the middle of the herd for defense according to trackways at Davenport Ranch, Texas. Dinosaurs may have congregated in herds for defense, for migratory purposes, or to provide protection for their young. The interpretation of dinosaurs as gregarious has also extended to depicting carnivorous theropods as pack hunters working together to bring down large prey.^[36][37] However, this lifestyle is not found among the modern relatives of dinosaurs (crocodiles and other reptiles, and birds), and the taphonomic evidence suggesting pack hunting in such theropods as Deinonychus and Allosaurus can also be interpreted as the results of fatal disputes between feeding animals, as is seen in many modern diapsid predators.^[38]

Jack Horner's 1978 discovery of a Maiasaura ("good mother dinosaur") nesting ground in Montana demonstrated that parental care continued long after birth among the ornithopods.^[39] There is also evidence that other Cretaceous-era dinosaurs, like Patagonian titanosaurian sauropods (1997 discovery), had similar nesting behaviors,^[40] and that the animals congregated in huge nesting colonies like those of penguins. The Mongolian oviraptorid Citipati was discovered in a chicken-like brooding position in 1993, which may mean it was covered with an insulating layer of feathers that kept the eggs warm.^[41] Parental care is also implied by the fossilized remains of a grouping of Psittacosaurus consisting of one adult and 34 juveniles; in this case, the large number of juveniles may be due to communal nesting.^[42] Trackways have also confirmed parental behavior among sauropods and ornithopods from the Isle of Skye in northwestern Scotland.^[43] 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 of some dinosaurs, like the marginocephalians, theropods and lambeosaurines, may have been too fragile to be used for active defense, so they were likely used for sexual or aggressive displays, though little is known about dinosaur mating and territorialism. Head wounds from bites suggest that theropods, at least, engaged in active aggressive confrontations.^[44] The nature of dinosaur communication also remains enigmatic, and is an active area of research. For example, recent studies suggest that the hollow crests of the lambeosaurines may have functioned as resonance chambers used for a wide range of vocalizations.^[45][46]

From a behavioral standpoint, one of the most valuable dinosaur fossils was discovered in the Gobi Desert in 1971. It included a Velociraptor attacking a Protoceratops,^[47] providing evidence that dinosaurs did indeed attack each other.^[48] Additional evidence for attacking live prey is the partially-healed tail of an Edmontosaurus, a hadrosaurid dinosaur; the tail is damaged in such a way that shows the animal was bitten by a tyrannosaur but survived. cannibalism amongst some species of dinosaurs was confirmed by tooth marks found in Madagascar in 2003, involving the theropod Majungasaurus.^[49]

Based on current fossil evidence from dinosaurs such as Oryctodromeus, some herbivorous species seem to have led a partially fossorial (burrowing) lifestyle,^[50] and some bird-like species may have been arboreal (tree-climbing), most notably primitive dromaeosaurids such as Microraptor^[51] and the enigmatic scansoriopterygids.^[52] 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, 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,^[53] whether diplodocids could create sonic booms via whip-like tail snapping,^[54] and whether sauropods could float.^[55]

Areas of controversy

Physiology

Main article: Physiology of dinosaurs

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 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 ectothermic creatures: "terrible lizards" as their name suggests. This supposed cold-bloodedness implied that dinosaurs were relatively slow, sluggish organisms, comparable to modern reptiles, which need external sources of heat in order to regulate their body temperature. Dinosaur ectothermy remained a prevalent view until Robert T. "Bob" Bakker, 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 (perhaps aided by the animals' bulk). Evidence of endothermism in dinosaurs includes the discovery of polar dinosaurs in Australia 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 that are typical of endotherms within dinosaur bone. Skeletal structures suggest that theropods and other dinosaurs had active lifestyles better suited to an endothermic cardiovascular system, while sauropods exhibit fewer endothermic characteristics. It is certainly possible that some dinosaurs were endothermic while others were not. Scientific debate over the specifics continues.^[56]

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 to average birds or mammals, which expend energy to elevate body temperature above that of the environment. Small birds and mammals also possess insulation, such as fat, fur, or feathers, 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 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 are hairless, and 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, 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 vast number of dog- and goat-sized dinosaur species which made up the bulk of the ecosystem during the Mesozoic Era.

Soft tissue and DNA

One of the best examples of soft tissue impressions in a fossil dinosaur was discovered in Petraroia, Italy. The discovery was reported in 1998, and described the specimen of a small, very young coelurosaur, Scipionyx samniticus. The fossil includes portions of the intestines, colon, liver, muscles, and windpipe of this immature dinosaur.^[26]

In the March 2005 issue of Science, Dr. Mary Higby Schweitzer and her team announced the discovery of flexible material resembling actual soft tissue inside a 68-million-year-old Tyrannosaurus rex leg bone from the Hell Creek Formation in Montana. After recovery, the tissue was rehydrated by the science team.^[27]

When the fossilized bone was treated over several weeks to remove mineral content from the fossilized bone marrow cavity (a process called demineralization), Schweitzer found evidence of intact structures such as blood vessels, bone matrix, and connective tissue (bone fibers). Scrutiny under the microscope further revealed that the putative dinosaur soft tissue had retained fine structures (microstructures) even at the cellular level. The exact nature and composition of this material, and the implications of Dr. Schweitzer's discovery, are not yet clear; study and interpretation of the material is ongoing.^[27]

The successful extraction of ancient DNA from dinosaur fossils has been reported on two separate occasions, but upon further inspection and peer review, neither of these reports could be confirmed.^[57] However, a functional visual peptide of a theoretical dinosaur has been inferred using analytical phylogenetic reconstruction methods on gene sequences of related modern species such as reptiles and birds.^[58] In addition, several proteins have putatively been detected in dinosaur fossils,^[59] including hemoglobin.^[60]

Even if dinosaur DNA could be reconstructed, it would be exceedingly difficult to clone and "grow" dinosaurs using current technology since no closely related species exist to provide zygotes or a suitable environment for embryonic development.

Feathered dinosaurs and the origin of birds

Main article: Feathered dinosaurs

Main article: Origin of birds

Birds and non-avian dinosaurs share many features. Birds share over a hundred distinct anatomical features with theropod dinosaurs, which are generally accepted to have been their closest ancient relatives.^[61]

Feathers

Archaeopteryx, the first good example of a "feathered dinosaur", was discovered in 1861. The initial specimen was found in the Solnhofen limestone in southern Germany, which is a lagerstätte, a rare and remarkable geological formation known for its superbly detailed fossils. Archaeopteryx is a transitional fossil, with features clearly intermediate between those of modern reptiles and birds. Brought to light just two years after Darwin's seminal The Origin of Species, its discovery spurred the nascent debate between proponents of evolutionary biology and creationism. This early bird is so dinosaur-like that, without a clear impression of feathers in the surrounding rock, at least one specimen was mistaken for Compsognathus.^[62]

Since the 1990s, a number of additional feathered dinosaurs have been found, providing even stronger evidence of the close relationship between dinosaurs and modern birds. Most of these specimens were unearthed in Liaoning province, northeastern China, which was part of an island continent during the Cretaceous period. Though feathers have been found only in the lagerstätte of the Yixian Formation and a few other places, 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 due to the fact that delicate features like skin and feathers are not often preserved by fossilization and thus are absent from the fossil record.

A recent development in the debate centers around the discovery of impressions of "protofeathers" surrounding many dinosaur fossils. Said protofeathers suggest that the tyrannosauroids may have been feathered.^[63] However, others claim that these protofeathers are simply the result of the decomposition of collagenous fiber that underlaid the dinosaurs' integument.^[64]

The feathered dinosaurs discovered so far include Beipiaosaurus, Caudipteryx, Dilong, Microraptor, Protarchaeopteryx, Shuvuuia, Sinornithosaurus, Sinosauropteryx, and Jinfengopteryx. Dinosaur-like birds like Confuciusornis, which are anatomically closer to modern avians, have also been discovered. All of these specimens come from the same formation in northern China. The dromaeosauridae family in particular seems to have been heavily feathered, and at least one dromaeosaurid, Cryptovolans, may have been capable of flight.

Skeleton

Because feathers are often associated with birds, feathered dinosaurs are often touted as the missing link between birds and dinosaurs. However, the multiple skeletal features also shared by the two groups represent the more important link for paleontologists. Furthermore, it is increasingly clear that the relationship between birds and dinosaurs, and the evolution of flight, are more complex topics than previously realized. For example, while it was once believed that birds evolved from dinosaurs in one linear progression, some scientists, most notably Gregory S. Paul, conclude that dinosaurs such as the dromaeosaurs may have evolved from birds, losing the power of flight while keeping their feathers in a manner similar to the modern ostrich and other ratites.

Comparison of bird and dinosaur skeletons, as well as cladistic analysis, strengthens the case for the link, particularly for a branch of theropods called maniraptors. Skeletal similarities include the neck, pubis, wrist (semi-lunate carpal), arm and pectoral girdle, shoulder blade, clavicle and breast bone.

Reproductive biology

A discovery of features in a Tyrannosaurus rex skeleton recently provided even more evidence that dinosaurs and birds evolved from a common ancestor 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 in their limbs. This medullary bone, which is rich in calcium, forms a layer inside the hard outer bone that 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.^[65]

A dinosaur embryo (pertaining to the prosauropod Massospondylus) was found without teeth, indicating that some parental care was required to feed the young dinosaur.^[66] It is also possible that the adult dinosaurs regurgitated into a young dinosaur's mouth to provide sustenance, a behavior that is also characteristic of numerous modern bird species.

Lungs

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. The lungs of theropod dinosaurs (carnivores that walked on two legs and had birdlike feet) likely pumped air into hollow sacs in their skeletons, as is the case in birds. "What was once formally considered unique to birds was present in some form in the ancestors of birds", O'Connor said. The study was funded in part by the National Science Foundation.^[67]

Heart and sleeping posture

Modern computerized tomography (CT) scans of a dinosaur chest cavity (conducted in 2000) found the apparent remnants of complex four-chambered hearts, much like those found in today's mammals and birds.^[68] The idea is controversial within the scientific community, coming under-fire for bad anatomical science^[69] or simply wishful thinking.^[70] A recently discovered troodont fossil demonstrates that the dinosaurs slept like certain modern birds, with their heads tucked under their arms.^[71] This behavior, which may have helped to keep the head warm, is also characteristic of modern birds.

Gizzard

Another piece of evidence that birds and dinosaurs are closely related is the use of gizzard stones. These stones are swallowed by animals to aid digestion and break down food and hard fibres once they enter the stomach. When found in association with fossils, gizzard stones are called gastroliths.^[72]

Extinction

Main article: Cretaceous–Tertiary extinction event

Main article: K–T boundary

Non-avian dinosaurs suddenly became extinct approximately 65 million years ago. Many other groups of animals also became extinct at this time, including ammonites (nautilus-like mollusks), mosasaurs, plesiosaurs, pterosaurs, herbivorous turtles and crocodiles, most birds, and many groups of mammals.^[7] This mass extinction is known as the Cretaceous–Tertiary 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 general 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 dinosaur era, there were no polar ice caps, and sea levels are estimated to have been from 100 meters (328 ft) to 250 meters (820 ft) higher than they are today. The planet's temperature was also much more uniform, with only +25 °C (77 °F) separating average polar temperatures from those at the equator. On average, atmospheric temperatures were also much warmer; the poles, for example, were +50 °C (122 °F) warmer than today.^[73][74]

The atmosphere's composition during the dinosaur era 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.^[7]

Impact event

The asteroid collision theory, which was first proposed by Walter Alvarez in the late 1970s, links the extinction event at the end of the Cretaceous period to a bolide impact approximately 65.5 million years ago. Alvarez proposed that a sudden increase in iridium 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 5 kilometers (3 mi) to 15 kilometers (9 mi) wide bolide hit in the vicinity of the Yucatán Peninsula, creating the 170 kilometers (106 mi) wide Chicxulub Crater and triggering the mass extinction. Scientists are not certain whether dinosaurs were thriving or declining before the impact event. 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.

Although the speed of extinction cannot be deduced from the fossil record alone, various models suggest that the extinction was extremely rapid. 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).

In September of 2007, U.S. researchers led by William Bottke of the Southwest Research Institute in Boulder, Colorado, and Czech scientists used computer simulations to identify the probable source of the Chicxulub impact. They calculated a 90% probability that a giant asteroid named Baptistina, approximately 160 kilometers (99 mi) in diameter, orbiting in the asteroid belt which lies between Mars and Jupiter, was struck by a smaller unnamed asteroid about 55 kilometers (34 mi) in diameter about 160 million years ago. The impact shattered Baptistina, creating a cluster which still exists today as the Baptistina family. Calculations indicate that some of the fragments were sent hurtling into earth-crossing orbits, one of which was the 10 kilometers (6 mi) wide meteorite which struck Mexico's Yucatan peninsula 65 million years ago, creating the Chicxulub crater (175 kilometers (109 mi)).

While similar to Alvarez's impact theory (which involved a single asteroid or comet), this theory proposes that "passages of the solar companion star Nemesis through the Oort comet cloud would trigger comet showers."^[75] One or more of these objects 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.^[75]

Deccan Traps

Main article: Deccan Traps

Before 2000, arguments that the Deccan Traps flood basalts caused the extinction were usually linked to the view that the extinction was gradual, as the flood basalt events were thought to have started around 68 mya and lasted for over 2 million years. However, there is evidence that two-thirds of the Deccan Traps were created in 1 million years about 65.5 mya, so these eruptions would have caused a fairly rapid extinction, possibly a period of thousands of years, but still a longer period than what would be expected from a single impact event.^[76][77]

The Deccan Traps could have caused extinction through several mechanisms, including the release of dust and sulphuric aerosols into the air which might have blocked sunlight and thereby reducing photosynthesis in plants. In addition, Deccan Trap volcanism might have resulted in carbon dioxide emissions which would have increased the greenhouse effect when the dust and aerosols cleared from the atmosphere.^[77] Before the mass extinction of the dinosaurs, the release of volcanic gasses during the formation of the Deccan traps "contributed to an apparently massive global warming. Some data point to an average rise in temperature of +8 °C (46.4 °F) in the last half million years before the impact [at Chicxulub]."^[78]

In the years when the Deccan Traps theory was linked to a slower extinction, Luis Alvarez (who died in 1988) replied that paleontologists were being misled by sparse data. While his assertion was not initially well-received, later intensive field studies of fossil beds lent weight to his claim. Eventually, most paleontologists began to accept the idea that the mass extinctions at the end of the Cretaceous were largely or at least partly due to a massive Earth impact. However, even Walter Alvarez has acknowledged that there were other major changes on Earth even before the impact, such as a drop in sea level and massive volcanic eruptions that produced the Indian Deccan Traps, and these may have contributed to the extinctions.^[