Dinosaur
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, although the exact origin and timing of the evolution of dinosaurs is a subject of active research. They became the dominant terrestrial vertebrates after the Triassic–Jurassic extinction event 201.3 mya and 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 11,000 living species, are among the most diverse groups 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 and fossil remains. Through most 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 are generally small due to the constraints of flight, many non-avian dinosaurs 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 first dinosaur fossils were recognized in the early 19th century, with the name "dinosaur" being coined by Sir Richard Owen in 1842 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 the Jurassic Park franchise. 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 of Triceratops and modern birds, 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 cover the same known genera: Dinosauria = Ornithischia + Saurischia. This includes major groups such as ankylosaurians, stegosaurians, ceratopsians, pachycephalosaurians, ornithopods, theropods, and sauropodomorphs.Birds are the sole surviving dinosaurs. In traditional taxonomy, birds were considered a separate class that had evolved from dinosaurs. However, most contemporary paleontologists reject the traditional style of classification based on anatomical similarity, in favor of phylogenetic taxonomy based on deduced ancestry, in which each group is defined as all descendants of a given founding genus. Birds belong to the dinosaur subgroup Maniraptora, which are coelurosaurs, which are theropods, which are saurischians.
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. This would cause sauropods and kin to fall outside traditional dinosaurs, so they re-defined Dinosauria 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. They also resurrected the clade Ornithoscelida to refer to the group containing Ornithischia and Theropoda.
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, except Dimetrodon. 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. Dinosaurs have always been recognized as an extremely varied group: over 900 non-avian dinosaur genera have been confidently identified with 1124 species. Estimates put the total number of dinosaur genera preserved in the fossil record at 1850, nearly 75% still undiscovered, and the number that ever existed at 3,400. A 2016 estimate put the number of dinosaur species living in the Mesozoic at 1,543–2,468, compared to the number of modern-day birds at 10,806 species.
Extinct dinosaurs, as well as modern birds, include genera that are herbivorous and others carnivorous, including seed-eaters, fish-eaters, insectivores, and omnivores. While dinosaurs were ancestrally bipedal, some evolved into quadrupeds, and others, such as Anchisaurus and Iguanodon, could walk 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 the best-known genera are remarkable for their 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 the Early Jurassic epoch at latest. Modern birds inhabit most available habitats, from terrestrial to marine, and there is evidence that some non-avian dinosaurs 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 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 neck vertebrae behind the atlas and axis, the first two neck vertebrae
- Apex of a deltopectoral crest located at or more than 30% down the length of the humerus
- Radius, a lower arm bone, shorter than 80% of humerus length
- Fourth trochanter on the femur 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 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
- Cnemial crest on the tibia arcs anterolaterally
- Distinct proximodistally oriented ridge present on the posterior face of the distal end of the tibia
- Concave articular surface for the fibula of the calcaneum
File:Sprawling and erect hip joints - horizontal.svg|thumb|Hip joints and hindlimb postures of: typical reptiles, dinosaurs and mammals, and rauisuchians
A variety of other skeletal features are shared by dinosaurs. However, because they either are common to other groups of archosaurs or were not present in all early dinosaurs, these features are not synapomorphies. For example, as diapsids, dinosaurs ancestrally had two pairs of Infratemporal fenestrae, 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 ; and a perforate acetabulum, or hip socket, with a hole at the center of its inside surface. 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 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.