Evolution of dinosaurs

Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is the subject of active research. They became the dominant terrestrial vertebrates after the Triassic–Jurassic extinction event 201.3 mya; their dominance continued throughout the Jurassic and Cretaceous periods. The fossil record shows that birds are feathered dinosaurs, having evolved from earlier theropods during the Late Jurassic epoch, and are the only dinosaur lineage known to have survived the Cretaceous–Paleogene extinction event approximately 66 mya. Dinosaurs can therefore be divided into avian dinosaurs—birds—and the extinct non-avian dinosaurs, which are all dinosaurs other than birds.

Dinosaurs are varied from taxonomic, morphological and ecological standpoints. Birds, at over 10,700 living species, are among the most diverse group of vertebrates. Using fossil evidence, paleontologists have identified over 900 distinct genera and more than 1,000 different species of non-avian dinosaurs. Dinosaurs are represented on every continent by both extant species (birds) and fossil remains. Through the first half of the 20th century, before birds were recognized as dinosaurs, most of the scientific community believed dinosaurs to have been sluggish and cold-blooded. Most research conducted since the 1970s, however, has indicated that dinosaurs were active animals with elevated metabolisms and numerous adaptations for social interaction. Some were herbivorous, others carnivorous. Evidence suggests that all dinosaurs were egg-laying, and that nest-building was a trait shared by many dinosaurs, both avian and non-avian.

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. While the dinosaurs' modern-day surviving avian lineage (birds) are generally small due to the constraints of flight, many prehistoric dinosaurs (non-avian and avian) were large-bodied—the largest sauropod dinosaurs are estimated to have reached lengths of and heights of and were the largest land animals of all time. The misconception that non-avian dinosaurs were uniformly gigantic is based in part on preservation bias, as large, sturdy bones are more likely to last until they are fossilized. Many dinosaurs were quite small, some measuring about in length.

The first dinosaur fossils were recognized in the early 19th century, with the name "dinosaur" (meaning "terrible lizard") being coined by Sir Richard Owen in 1841 to refer to these "great fossil lizards". Since then, mounted fossil dinosaur skeletons have been major attractions at museums worldwide, and dinosaurs have become an enduring part of popular culture. The large sizes of some dinosaurs, as well as their seemingly monstrous and fantastic nature, have ensured their regular appearance in best-selling books and films, such as '' Jurassic Park''. Persistent public enthusiasm for the animals has resulted in significant funding for dinosaur science, and new discoveries are regularly covered by the media.

Definition

Under phylogenetic nomenclature, dinosaurs are usually defined as the group consisting of the most recent common ancestor (MRCA) of ''Triceratops'' and modern birds (Neornithes), and all its descendants. 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. Both definitions result in the same set of animals being defined as dinosaurs: "Dinosauria = Ornithischia + Saurischia". This definition includes major groups such as ankylosaurians (armored herbivorous quadrupeds), stegosaurians (plated herbivorous quadrupeds), ceratopsians (bipedal or quadrupedal herbivores with neck frills), pachycephalosaurians (bipedal herbivores with thick skulls), ornithopods (bipedal or quadrupedal herbivores including " duck-bills"), theropods (mostly bipedal carnivores and birds), and sauropodomorphs (mostly large herbivorous quadrupeds with long necks and tails).

Birds are now recognized as being the sole surviving lineage of theropod dinosaurs. In traditional taxonomy, birds were considered a separate class that had evolved from dinosaurs, a distinct superorder. However, a majority of contemporary paleontologists concerned with dinosaurs reject the traditional style of classification in favor of phylogenetic taxonomy; this approach requires that, for a group to be natural, all descendants of members of the group must be included in the group as well. Birds are thus considered to be dinosaurs and dinosaurs are, therefore, not extinct. Birds are classified as belonging to the subgroup Maniraptora, which are coelurosaurs, which are theropods, which are saurischians, which are dinosaurs.

Research by Matthew G. Baron, David B. Norman, and Paul M. Barrett in 2017 suggested a radical revision of dinosaurian systematics. Phylogenetic analysis by Baron ''et al.'' recovered the Ornithischia as being closer to the Theropoda than the Sauropodomorpha, as opposed to the traditional union of theropods with sauropodomorphs. They resurrected the clade Ornithoscelida to refer to the group containing Ornithischia and Theropoda. Dinosauria itself was re-defined as the last common ancestor of ''Triceratops horridus'', ''Passer domesticus'' and ''Diplodocus carnegii'', and all of its descendants, to ensure that sauropods and kin remain included as dinosaurs. "A version of this article appears in print on March 28, 2017, on Page D6 of the New York edition with the headline: Shaking Up the Dinosaur Family Tree." "This file contains Supplementary Text and Data, Supplementary Tables 1-3 and additional references."
Supplementary Information
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General description

Using one of the above definitions, dinosaurs can be generally described as archosaurs with hind limbs held erect beneath the body. Other prehistoric animals, including pterosaurs, mosasaurs, ichthyosaurs, plesiosaurs, and '' Dimetrodon'', while often popularly conceived of as dinosaurs, are not taxonomically classified as dinosaurs. Pterosaurs are distantly related to dinosaurs, being members of the clade Ornithodira. The other groups mentioned are, like dinosaurs and pterosaurs, members of Sauropsida (the reptile and bird clade), except ''Dimetrodon'' (which is a synapsid). None of them had the erect hind limb posture characteristic of true dinosaurs.

Dinosaurs were the dominant terrestrial vertebrates of the Mesozoic Era, especially the Jurassic and Cretaceous periods. Other groups of animals were restricted in size and niches; mammals, for example, rarely exceeded the size of a domestic cat, and were generally rodent-sized carnivores of small prey. They have always been recognized as an extremely varied group of animals; over 900 non-avian dinosaur genera have been identified with certainty as of 2018, and the total number of genera preserved in the fossil record has been estimated at around 1850, nearly 75% of which remain to be discovered, and 1124 species by 2016. A 1995 study predicted that about 3,400 dinosaur genera ever existed, including many that would not have been preserved in the fossil record.

In 2016, the estimated number of dinosaur species that existed in the Mesozoic was 1,543–2,468. In 2021, the number of modern-day birds (avian dinosaurs) was estimated to be at 10,806 species. Some are herbivorous, others carnivorous, including seed-eaters, fish-eaters, insectivores, and omnivores. While dinosaurs were ancestrally bipedal (as are all modern birds), some prehistoric species were quadrupeds, and others, such as ''Anchisaurus'' and ''Iguanodon'', could walk just as easily on two or four legs. Cranial modifications like horns and crests are common dinosaurian traits, and some extinct species had bony armor. Although known for large size, many Mesozoic dinosaurs were human-sized or smaller, and modern birds are generally small in size. Dinosaurs today inhabit every continent, and fossils show that they had achieved global distribution by at least the Early Jurassic epoch. Modern birds inhabit most available habitats, from terrestrial to marine, and there is evidence that some non-avian dinosaurs (such as ''Microraptor'') could fly or at least glide, and others, such as spinosaurids, had semiaquatic habits.

Distinguishing anatomical features

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

A detailed assessment of archosaur interrelations by Sterling Nesbitt confirmed or found the following twelve unambiguous synapomorphies, some previously known:
* In the skull, a supratemporal fossa (excavation) is present in front of the supratemporal fenestra, the main opening in the rear skull roof
* Epipophyses, obliquely backward-pointing processes on the rear top corners of the anterior (front) neck vertebrae behind the atlas and axis, the first two neck vertebrae
* Apex of a deltopectoral crest (a projection on which the deltopectoral muscles attach) located at or more than 30% down the length of the humerus (upper arm bone)
* Radius, a lower arm bone, shorter than 80% of humerus length
* Fourth trochanter (projection where the caudofemoralis muscle attaches on the inner rear shaft) on the femur (thigh bone) is a sharp flange
* Fourth trochanter asymmetrical, with distal, lower, margin forming a steeper angle to the shaft
* On the astragalus and calcaneum, upper ankle bones, the proximal articular facet, the top connecting surface, for the fibula occupies less than 30% of the transverse width of the element
* Exoccipitals (bones at the back of the skull) do not meet along the midline on the floor of the endocranial cavity, the inner space of the braincase
* In the pelvis, the proximal articular surfaces of the ischium with the ilium and the pubis are separated by a large concave surface (on the upper side of the ischium a part of the open hip joint is located between the contacts with the pubic bone and the ilium)
* Cnemial crest on the tibia (protruding part of the top surface of the shinbone) arcs anterolaterally (curves to the front and the outer side)
* Distinct proximodistally oriented (vertical) ridge present on the posterior face of the distal end of the tibia (the rear surface of the lower end of the shinbone)
* Concave articular surface for the fibula of the calcaneum (the top surface of the calcaneum, where it touches the fibula, has a hollow profile)

Nesbitt found a number of further potential synapomorphies and discounted a number of synapomorphies previously suggested. Some of these are also present in silesaurids, which Nesbitt recovered as a sister group to Dinosauria, including a large anterior trochanter, metatarsals II and IV of subequal length, reduced contact between ischium and pubis, the presence of a cnemial crest on the tibia and of an ascending process on the astragalus, and many others.

A variety of other skeletal features are shared by dinosaurs. However, because they are either common to other groups of archosaurs or were not present in all early dinosaurs, these features are not considered to be synapomorphies. For example, as diapsids, dinosaurs ancestrally had two pairs of Infratemporal fenestrae (openings in the skull behind the eyes), and as members of the diapsid group Archosauria, had additional openings in the snout and lower jaw. 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''); and a perforate acetabulum, or hip socket, with a hole at the center of its inside surface (closed in ''Saturnalia tupiniquim'', for example). Another difficulty of determining distinctly dinosaurian features is that early dinosaurs and other archosaurs from the Late Triassic epoch are often poorly known and were similar in many ways; these animals have sometimes been misidentified in the literature.

Dinosaurs stand with their hind limbs erect in a manner similar to most modern mammals, but distinct from most other reptiles, whose limbs sprawl out to either side. This posture is due to the development of a laterally facing recess in the pelvis (usually an open socket) and a corresponding inwardly facing distinct head on the femur. Their erect posture enabled early dinosaurs to breathe easily while moving, which likely permitted stamina and activity levels that surpassed those of "sprawling" reptiles. Erect limbs probably also helped support the evolution of large size by reducing bending stresses on limbs. 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.

History of study

Pre-scientific history

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

Early dinosaur research

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

Between 1815 and 1824, the Rev William Buckland, the first Reader of Geology at the University of Oxford, collected more fossilized bones of ''Megalosaurus'' and became the first person to describe a non-avian dinosaur in a scientific journal. The second non-avian dinosaur genus to be identified, ''Iguanodon'', was discovered in 1822 by Mary Ann Mantell – the wife of English geologist Gideon Mantell. Gideon Mantell recognized similarities between his fossils and the bones of modern iguanas. He published his findings in 1825.

The study of these "great fossil lizards" soon became of great interest to European and American scientists, and in 1841 the English paleontologist Sir Richard Owen coined the term "dinosaur", using it to refer to the "distinct tribe or sub-order of Saurian Reptiles" that were then being recognized in England and around the world. The term is derived . Though the taxonomic name has often been interpreted as a reference to dinosaurs' teeth, claws, and other fearsome characteristics, Owen intended it to also evoke their size and majesty. Owen recognized that the remains that had been found so far, ''Iguanodon'', ''Megalosaurus'' and ''Hylaeosaurus'', shared a number of distinctive features, and so decided to present them as a distinct taxonomic group. With the backing of Prince Albert, the husband of Queen Victoria, Owen established the Natural History Museum, London, to display the national collection of dinosaur fossils and other biological and geological exhibits.

Discoveries in North America

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

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

"Dinosaur renaissance" and beyond

The field of dinosaur research has enjoyed a surge in activity that began in the 1970s and is ongoing. This was triggered, in part, by John Ostrom's discovery and 1969 description of ''Deinonychus'', an active predator that may have been warm-blooded, in marked contrast to the then-prevailing image of dinosaurs as sluggish and cold-blooded. Vertebrate paleontology has become a global science. Major new dinosaur discoveries have been made by paleontologists working in previously unexploited regions, including India, South America, Madagascar, Antarctica, and most significantly China (the well-preserved feathered dinosaurs in China have further consolidated the link between dinosaurs and their living descendants, modern birds). The widespread application of cladistics, which rigorously analyzes the relationships between biological organisms, has also proved tremendously useful in classifying dinosaurs. Cladistic analysis, among other modern techniques, helps to compensate for an often incomplete and fragmentary fossil record.

Soft tissue and DNA

One of the best examples of soft-tissue impressions in a fossil dinosaur was discovered in the Pietraroja Plattenkalk in southern Italy. The discovery was reported in 1998, and described the specimen of a small, juvenile coelurosaur, ''Scipionyx samniticus''. The fossil includes portions of the intestines, colon, liver, muscles, and windpipe of this dinosaur.

In the March 2005 issue of ''Science'', the paleontologist Mary Higby Schweitzer and her team announced the discovery of flexible material resembling actual soft tissue inside a 68-million-year-old ''Tyrannosaurus rex'' leg bone from the Hell Creek Formation in Montana. After recovery, the tissue was rehydrated by the science team. When the fossilized bone was treated over several weeks to remove mineral content from the fossilized bone-marrow cavity (a process called demineralization), Schweitzer found evidence of intact structures such as blood vessels, bone matrix, and connective tissue (bone fibers). Scrutiny under the microscope further revealed that the putative dinosaur soft tissue had retained fine structures (microstructures) even at the cellular level. The exact nature and composition of this material, and the implications of Schweitzer's discovery, are not yet clear.

In 2009, a team including Schweitzer announced that, using even more careful methodology, they had duplicated their results by finding similar soft tissue in a duck-billed dinosaur, ''Brachylophosaurus canadensis'', found in the Judith River Formation of Montana. This included even more detailed tissue, down to preserved bone cells that seem to have visible remnants of nuclei and what seem to be red blood cells. Among other materials found in the bone was collagen, as in the ''Tyrannosaurus'' bone. The type of collagen an animal has in its bones varies according to its DNA and, in both cases, this collagen was of the same type found in modern chickens and ostriches.

The extraction of ancient DNA from dinosaur fossils has been reported on two separate occasions; upon further inspection and peer review, however, neither of these reports could be confirmed. However, a functional peptide involved in the vision of a theoretical dinosaur has been inferred using analytical phylogenetic reconstruction methods on gene sequences of related modern species such as reptiles and birds. In addition, several proteins, including hemoglobin, have putatively been detected in dinosaur fossils.

In 2015, researchers reported finding structures similar to blood cells and collagen fibers, preserved in the bone fossils of six Cretaceous dinosaur specimens, which are approximately 75 million years old.

Evolutionary history

Origins and early evolution

Dinosaurs diverged from their archosaur ancestors during the Middle to Late Triassic epochs, roughly 20 million years after the devastating Permian–Triassic extinction event wiped out an estimated 96% of all marine species and 70% of terrestrial vertebrate species approximately 252 million years ago. Radiometric dating of the Ischigualasto Formation of Argentina where the early dinosaur genus ''Eoraptor'' was found date it as 231.4 million years old. ''Eoraptor'' is thought to resemble the common ancestor of all dinosaurs; if this is true, its traits suggest that the first dinosaurs were small, bipedal predators. The discovery of primitive, dinosaur-like ornithodirans such as ''Lagosuchus'' and ''Lagerpeton'' in Argentina in the Carnian epoch of the Triassic, around 233 million years ago, supports this view; analysis of recovered fossils suggests that these animals were indeed small, bipedal predators. Dinosaurs may have appeared as early as the Anisian epoch of the Triassic, 245 million years ago, as evidenced by remains of the genus ''Nyasasaurus'' from that period. However, its known fossils are too fragmentary to tell if it was a dinosaur or only a close relative. Paleontologist Max C. Langer ''et al.'' (2018) determined that '' Staurikosaurus'' from the Santa Maria Formation dates to 233.23 million years ago, making it older in geologic age than ''Eoraptor''.

When dinosaurs appeared, they were not the dominant terrestrial animals. The terrestrial habitats were occupied by various types of archosauromorphs and therapsids, like cynodonts and rhynchosaurs. Their main competitors were the pseudosuchians, such as aetosaurs, ornithosuchids and rauisuchians, which were more successful than the dinosaurs. Most of these other animals became extinct in the Triassic, in one of two events. First, at about 215 million years ago, a variety of basal archosauromorphs, including the protorosaurs, became extinct. This was followed by the Triassic–Jurassic extinction event (about 201 million years ago), that saw the end of most of the other groups of early archosaurs, like aetosaurs, ornithosuchids, phytosaurs, and rauisuchians. Rhynchosaurs and dicynodonts survived (at least in some areas) at least as late as early –mid Norian and late Norian or earliest Rhaetian stages, respectively, and the exact date of their extinction is uncertain. These losses left behind a land fauna of crocodylomorphs, dinosaurs, mammals, pterosaurians, and turtles. 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. Also notably, there was a heightened rate of extinction during the Carnian pluvial event.

Evolution and paleobiogeography

Dinosaur evolution after the Triassic followed changes in vegetation and the location of continents. In the Late Triassic and Early Jurassic, the continents were connected as the single landmass Pangaea, and there was a worldwide dinosaur fauna mostly composed of coelophysoid carnivores and early sauropodomorph herbivores. 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. The general homogeneity of dinosaurian faunas continued into the Middle and Late Jurassic, where most localities had predators consisting of ceratosaurians, megalosauroids, and allosauroids, and herbivores consisting of stegosaurian ornithischians and large sauropods. Examples of this include the Morrison Formation of North America and Tendaguru Beds of Tanzania. Dinosaurs in China show some differences, with specialized metriacanthosaurid theropods and unusual, long-necked sauropods like ''Mamenchisaurus''. Ankylosaurians and ornithopods were also becoming more common, but primitive sauropodomorphs had become extinct. Conifers and pteridophytes were the most common plants. Sauropods, like earlier sauropodomorphs, were not oral processors, but ornithischians were evolving various means of dealing with food in the mouth, including potential cheek-like organs to keep food in the mouth, and jaw motions to grind food. Another notable evolutionary event of the Jurassic was the appearance of true birds, descended from maniraptoran coelurosaurians.

By the Early Cretaceous and the ongoing breakup of Pangaea, dinosaurs were becoming strongly differentiated by landmass. The earliest part of this time saw the spread of ankylosaurians, iguanodontians, and 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. The stegosaurians appear to have gone extinct at some point in the late Early Cretaceous or early Late Cretaceous. A major change in the Early Cretaceous, which would be amplified in the Late Cretaceous, was the evolution of flowering plants. At the same time, several groups of dinosaurian herbivores evolved more sophisticated ways to orally process food. Ceratopsians developed a method of slicing with teeth stacked on each other in batteries, and iguanodontians refined a method of grinding with dental batteries, taken to its extreme in hadrosaurids. Some sauropods also evolved tooth batteries, best exemplified by the rebbachisaurid ''Nigersaurus''.

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

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

The surviving lineages of neornithine birds, including the ancestors of modern ratites, ducks and chickens, and a variety of waterbirds, diversified rapidly at the beginning of the Paleogene period, entering ecological niches left vacant by the extinction of Mesozoic dinosaur groups such as the arboreal enantiornithines, aquatic hesperornithines, and even the larger terrestrial theropods (in the form of ''Gastornis'', eogruiids, bathornithids, ratites, geranoidids, mihirungs, and "terror birds"). It is often stated that mammals out-competed the neornithines for dominance of most terrestrial niches but many of these groups co-existed with rich mammalian faunas for most of the Cenozoic Era. Terror birds and bathornithids occupied carnivorous guilds alongside predatory mammals, and ratites are still fairly successful as mid-sized herbivores; eogruiids similarly lasted from the Eocene to Pliocene

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