Introduction on Dinosaur

Areas of debate

Physiology

    A vigorous debate on the subject of temperature regulation in dinosaurs has been ongoing since the 1960s. Originally, scientists broadly disagreed as to whether dinosaurs were capable of regulating their body temperatures at all. More recently, dinosaur endothermy 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.

    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 period.

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.

    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 the Liaoning province in 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. However, others claim that these protofeathers are simply the result of the decomposition of collagenous fiber that underlaid the dinosaurs' integument.

    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.

    A dinosaur embryo was found without teeth, suggesting that some parental care was required to feed the young dinosaur. 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.

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. The idea is controversial within the scientific community, coming under-fire for bad anatomical science or simply wishful thinking. A recently discovered troodont fossil demonstrates that the dinosaurs slept like certain modern birds, with their heads tucked under their arms. 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. Because a particular stone could have been swallowed at one location before being carried to another during migration, paleontologists sometimes use the stones found in dinosaur stomachs to establish possible migration routes.

Evidence for Cenozoic dinosaurs

    In 2002, paleontologists Zielinski and Budahn reported the discovery of a single hadrosaur leg bone fossil in the San Juan Basin, New Mexico. The formation in which the bone was discovered has been dated to the early Paleocene epoch approximately 64.5 million years ago. If the bone was not re-deposited into that stratum by weathering action, it would provide evidence that some dinosaur populations may have survived at least a half million years into the Cenozoic 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.

    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.

    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 artifacts is ongoing.

    The successful extraction of ancient DNA from dinosaur fossils has been reported on two separate occasions, but upon further inspection and peer review, neither of these reports could be confirmed. 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. In addition, several proteins have putatively been detected in dinosaur fossils, including hemoglobin.

    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.

Extinction theories

    The sudden mass extinction of non-avian dinosaurs, an event that occurred approximately 65 million years ago, is one of the most intriguing mysteries in paleontology. 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. 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.

Asteroid collision

    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 5–15 km wide bolide hit in the vicinity of the Yucatán Peninsula, creating the 170 km-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).

Multiple collisions—the Oort cloud

    While similar to Alvarez's impact theory (which involved a single asteroid or comet), this theory proposes that a stream of comets was dislodged from the Oort cloud due to the gravitational disruption caused by a passing star. 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.

Environment changes

    At the peak of the dinosaur era, there were no polar ice caps, and sea levels are estimated to have been from 100 to 250 metres (330 to 820 feet) higher than they are today. The planet's temperature was also much more uniform, with only 25 degrees Celsius separating average polar temperatures from those at the equator. On average, atmospheric temperatures were also much warmer; the poles, for example, were 50 °C warmer than today.

    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.

History of discovery

    Dinosaur fossils have been known for millennia, although their true nature was not recognized. The Chinese, whose own word for dinosaur is konglong (恐龍, or "terrible dragon"), considered them to be dragon bones and documented them as such. For example, Hua Yang Guo Zhi, a book written by Zhang Qu during the Western Jin Dynasty, reported the discovery of dragon bones at Wucheng in Sichuan Province.^[59] In Europe, dinosaur fossils were generally believed to be the remains of giants and other creatures killed by the Great Flood.

    Megalosaurus was the first dinosaur to be formally described, in 1677, when part of a bone was recovered from a limestone quarry at Cornwell near Oxford, England. This bone fragment was identified correctly as the lower extremity of the femur of an animal larger than anything living in modern times. The second dinosaur genus to be identified, Iguanodon, was discovered in 1822 by the English geologist Gideon Mantell, who recognized similarities between his fossils and the bones of modern iguanas. Two years later, the Rev William Buckland, a professor of geology at Oxford University, unearthed more fossilized bones of Megalosaurus and became the first person to describe dinosaurs in a scientific journal.

    The study of these "great fossil lizards" soon became of great interest to European and American scientists, and in 1842 the English paleontologist Richard Owen coined the term "dinosaur". He 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 of Saxe-Coburg-Gotha, the husband of Queen Victoria, Owen established the Natural History Museum in South Kensington, London, to display the national collection of dinosaur fossils and other biological and geological exhibits.

    In 1858, the first known American dinosaur was discovered, in marl 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 the first nearly complete dinosaur skeleton found and it was clearly a bipedal creature. This was a revolutionary discovery as, until that point, most scientists had believed dinosaurs walked on four feet, like other lizards. Foulke's discoveries sparked a wave of dinosaur mania in the United States.

    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. The feud probably originated when Marsh publicly pointed out that Cope's reconstruction of an Elasmosaurus skeleton was flawed; Cope had inadvertently placed the plesiosaur's head at what should have been the animal's tail end. The fight between the two scientists lasted for over 30 years, ending in 1897 when Cope died after spending his entire fortune on the dinosaur hunt. Marsh 'won' the contest primarily because he was better funded through a relationship with the US Geological Survey. Unfortunately, many valuable dinosaur specimens were damaged or destroyed due to the pair's rough methods; for example, their diggers often used dynamite to unearth bones (a method modern paleontologists would find appalling). Despite their unrefined methods, the contributions of Cope and Marsh to paleontology were vast; Marsh unearthed 86 new species of dinosaur and Cope discovered 56, for a total of 142 new species. Cope's collection is now at the American Museum of Natural History in New York, while Marsh's is on display at the Peabody Museum of Natural History at Yale University.

    Since 1897, the search for dinosaur fossils has extended to every continent, including Antarctica. The first Antarctic dinosaur to be discovered, the ankylosaurid Antarctopelta oliveroi, was found on Ross Island in 1986, although it was 1994 before an Antarctic species, the theropod Cryolophosaurus ellioti, was formally named and described in a scientific journal.

    Current dinosaur "hot spots" include southern South America (especially Argentina) and China. China in particular has produced many exceptional feathered dinosaur specimens due to the unique geology of its dinosaur beds, as well as an ancient arid climate particularly conducive to fossilization.

Dinosaurs in culture

    By human standards, dinosaurs were creatures of fantastic appearance and often enormous size. As such, they have captured the public imagination and become an enduring part of human culture. Only three decades after the first scientific descriptions of dinosaur remains, the famous dinosaur sculptures were erected in Crystal Palace Park in London. These sculptures excited the public so strongly that smaller replicas were sold, one of the first examples of tie-in merchandising. Since Crystal Palace, dinosaur exhibitions have opened at parks and museums around the world, both catering to, and reinforcing, the public interest. Dinosaur popularity has long had a reciprocal effect on dinosaur science, as well. The competition between museums for public attention led directly to the Bone Wars waged between Marsh and Cope, each striving to return with more spectacular fossil remains than the other, and the resulting contribution to dinosaur science was enormous.

    Dinosaurs hold an integral place in modern culture. The word "dinosaur" itself has entered the English lexicon as an expression describing anything that is impractically large, slow-moving, or obsolete, bound for extinction. The public preoccupation with dinosaurs led to their inevitable entrance into worldwide popular culture. Beginning with a passing mention of Megalosaurus in the first paragraph of Charles Dickens' Bleak House in 1852, dinosaurs have been featured in a broad array of fictional works. Sir Arthur Conan Doyle's 1912 book The Lost World, the iconic 1933 film King Kong, the 1954 introduction of Godzilla and its many subsequent sequels, the best-selling 1990 novel Jurassic Park by Michael Crichton and its 1993 film version, briefly the highest-grossing film of all time, are just a few prominent examples of the long tradition of dinosaurs in fiction. Non-fiction authors, including some prominent paleontologists, have also sought to take advantage of dinosaur popularity, especially among children, to educate readers about dinosaurs in particular and science in general. Dinosaurs are also ubiquitous in advertising, with numerous companies seeking to utilize dinosaurs to sell their own products or to characterize their rivals as slow-moving or obsolete.

Religious views

    Various religious groups have views about dinosaurs that differ from those held by scientists, usually due to conflicts with creation stories in their scriptures. However, the scientific community does not accept these religiously-inspired interpretations of dinosaurs.