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The history of paleontology traces the effort to understand the history of life on Earth by studying the fossil record left behind by living organisms. Paleontology is a field of biology but its development has been closely tied to geology and the effort to understand the history of Earth itself. In Europe the systematic study of fossils emerged as an integral part of the changes in natural philosophy that occurred during the Age of reason. The nature of fossils and their relationship to life in the past became better understood during the 17th and 18th centuries, and at the end of the 18th century the work of Georges Cuvier led to the emergence of paleontology, in association with comparative anatomy, as a scientific discipline. The expanding knowledge of the fossil record also played an increasing role in the development of geology, particularly stratigraphy.

The first half of the 19th century saw a rapid increase in knowledge about the past history of life on Earth and progress towards definition of the geologic time scale. This made it increasingly obvious that there had been some kind of successive order to the development of life, and that would contribute to early theories of the transmutation of species.^[1] After Charles Darwin published Origin of Species in 1859, much of the focus of paleontology shifted to understanding evolutionary paths, including human evolution, and evolutionary theory.^[1]

The last half of the 19th century saw a tremendous expansion in paleontological activity, especially in North America. The trend continued in the 20th century with additional regions of the Earth being opened to systematic fossil collection, as demonstrated by a series of important discoveries in China. The last half of the 20th century saw a renewed interest in mass extinctions and their role in the evolution of life on Earth.^[2]

Prior to the 17th century

As early as the 6th century BC, Xenophanes of Colophon recognized that some fossil shells were remains of shellfish, which he used to argue that what was at the time dry land was once under the sea.^[3] Leonardo da Vinci (1452-1519), in an unpublished notebook, also concluded that some fossil sea shells were the remains of shellfish. However, in both cases, the fossils were complete remains of shellfish species that closely resembled living species, and were therefore easy to classify.^[4]

Shen Kuo (Chinese: 沈括) (1031 - 1095) of the Song Dynasty used the evidence of marine fossils found in the Taihang Mountains to infer the existence of geological processes of geomorphology and shifting of seashores over time.^[5] Using his observation of preserved petrified bamboos found underground in Yan'an, Shanbei region, Shaanxi province, he argued for a theory of gradual climate change, since Shaanxi was (and still is) part of a dry climate zone that does not support a habitat for the growth of bamboos.^[6]

As a result of a new emphasis on observing, classifying and cataloging nature, 16th century natural philosophers in Europe began to establish extensive collections of fossil objects (as well as collections of plant and animal specimens), which were often stored in specially built cabinets to help organize them. Conrad Gesner published a 1565 work on fossils that contained one of the first detailed descriptions of such a collection. The collection belonged to a member of the extensive network of correspondents that Gesner drew on for his works. Such informal correspondence networks among natural philosophers and collectors became increasingly important during the course of the 16th century and were direct forerunners of the scientific societies that would begin to form in the 17th century. These collections and correspondence networks played an important role in the development of natural philosophy.^[7]

However, most 16th century Europeans did not recognize that fossils were the remains of living organisms. The etymology of the word fossil comes from the Latin for things having been dug up. As this indicates, the term was applied to wide variety of stone and stone-like objects without regard to whether they might have an organic origin. Sixteenth century writers such as Gesner and Georg Agricola were more interested in classifying such objects by their physical and mystical properties than they were in determining the objects' origins.^[8] Also the natural philosophy of the period encouraged alternative explanations for the origin of fossils. Both the Aristotelian and Neoplatonic schools of philosophy provided support for the idea that stony objects might grow within the earth to resemble living things. Neoplatonic philosophy maintained that there could be affinities between living and non-living objects that could cause one to resemble the other. The Aristotelian school maintained that the seeds of living organisms could enter the ground and generate objects resembling those organisms.^[9]

17th century

During the Age of Reason, fundamental changes in natural philosophy were reflected in the analysis of fossils. In 1665 Robert Hooke published Micrographia, an illustrated collection of his observations with a microscope. One of these observations was entitled Of Petrify'd wood, and other Petrify'd bodies, which included a comparison between petrified and ordinary wood. He concluded that petrified wood was ordinary wood that had been soaked with "water impregnated with stony and earthy particles". He then suggested that several kinds of fossil sea shells were formed from ordinary shells by a similar process. He argued against the prevalent view that such objects were "Stones form'd by some extraordinary Plastick virtue latent in the Earth itself".^[10] In 1667 Nicholas Steno wrote a paper comparing the teeth of the shark with the common fossil objects known as tongue stones. He concluded that the fossils must have been shark teeth. Steno then took an interest in the question of fossils, and to address some of the objections to their ancient organic origin he began sudying rock strata. The result of this work was published in 1669 as Forerunner to a Dissertation on a solid naturally enclosed in a solid. In this book, Steno drew a clear distinction between objects such as rock crystals that really were formed within rocks and those such as fossil shells and shark teeth that were formed outside of those rocks. Steno realized that certain kinds of rock had been formed by the successive deposition of horizontal layers of sediment and that fossils were the remains of living organisms that had become buried in that sediment. Steno who, like almost all 17th century natural philosophers, believed that the earth was only a few thousand years old, resorted to the Biblical flood as a possible explanation for fossils of marine organism that were found very far from the sea.^[11]

Despite the considerable influence of Forerunner, naturalists such as Martin Lister (1638-1712) and John Ray (1627-1705) continued to question the organic origin of some fossils. They were particularly concerned about objects such as fossil Ammonites, which Hooke had claimed were organic in origin, that did not resemble any known living species. This raised the possibility of extinction, which they found difficult to accept for philosophical and theological reasons.^[12]

18th century

In his 1778 work Epochs of Nature Georges Buffon referred to fossils, in particular the discovery of fossils of tropical species such as elephants and rhinoceros in northern Europe, as evidence for the theory that the earth had started out much warmer than it currently was and had been gradually cooling.

In 1796 Georges Cuvier presented a paper on living and fossil elephants comparing skeletal remains of Indian and African elephants to fossils of mammoths and of an animal he would later name mastodon utilizing comparative anatomy. He established for the first time that Indian and African elephants were different species, and that mammoths differed from both and must be extinct. He further concluded that the mastodon was another extinct species that also differed from Indian or African elephants, more so than mammoths. Cuvier’s ground-breaking work in paleontology and comparative anatomy lead to the widespread acceptance of extinction.^[13] It also lead Cuvier to advocate the geological theory of catastrophism to explain the succession of organisms revealed by the fossil record. He also pointed out that since mammoths and wooly rhinoceros were not the same species as the elephants and rhinoceros currently living in the tropics, their fossils could not be used as evidence for a cooling earth. Cuvier made another powerful demonstration of the power of comparative anatomy in paleontology when he presented a second paper in 1796 on a large fossil skeleton from Paraguay, which he named Megatherium and identified as a giant sloth by comparing its skull to those of two living species of tree sloth.

In a pioneering application of stratigraphy, William Smith, a surveyor and mining engineer, made extensive use of fossils to help correlate rock strata in different locations. He created the first geological map of England during the late 1790s and early 1800s. He established the principle of faunal succession, the idea that each strata of sedimentary rock would contain particular types of fossils, and that these would succeed one another in a predictable way even in widely separated geologic formations. At the same time, Cuvier and Alexandre Brongniart, an instructor at the Paris school of mine engineering, used similar methods in an influential study of the geology of the region around Paris.

19th century before Darwin

The age of reptiles

In 1808, Cuvier identified a fossil found in Maastricht as a giant marine reptile that he named Mosasaurus. He also identified, from a drawing, another fossil found in Bavaria as a flying reptile and named it Pterodactylus. He speculated that an age of reptiles had preceded the first mammals.^[14]

Cuvier's speculation would be supported by a series of finds that would be made in Great Britain over the course of the next two decades. Mary Anning, a professional fossil collector since age 11, collected the fossils of a number of marine reptiles from the Jurassic marine strata at Lyme Regis. These included the first ichthyosaur skeleton to be recognized as such, which was collected in 1811, and the first plesiosaur collected in 1821. Many of her discoveries would be described scientifically by the geologists William Conybeare, Henry De la Beche and William Buckland.^[15]

In 1824, Buckland found and described a lower jaw from Jurassic deposits from Stonesfield. He determined that the bone belonged to a carnivorous land-dwelling reptile he called Megalosaurus. That same year Gideon Mantell realized that some large teeth he had found in 1822, in Cretaceous rocks from Tilgate, belonged to a giant herbivorous land-dwelling reptile. He called it Iguanodon, because the teeth resembled those of an iguana. In 1832 Mantell would find, in Tilgate, a partial skeleton of an armoured reptile he would call Hylaeosaurus. In 1842 the English anatomist Richard Owen would create a new order of reptiles, that he called Dinosauria for Megalosaurus, Iguanodon and Hylaeosaurus.^[16]

This evidence that giant reptiles had lived on Earth in the past caused great excitement in scientific circles,^[17] and even among some segments of the general public.^[18] Buckland did describe the jaw of a small primitive mammal, Phascolotherium, that was found in the same strata as Megalosaurus. This discovery, known as the Stonesfield mammal, was a much discussed anomaly. Cuvier at first thought it was a marsupial, but Buckland later realized it was a primitive placental mammal. Due to its small size and primitive nature, Buckland did not believe it invalidated the overall pattern of a time named "the age of reptiles", when the largest and most conspicuous animals had been reptiles rather than mammals.^[19]

Paleobotany

In 1828 Alexandre Brongniart's son, the botanist Adolphe Brongniart, published the introduction to a longer work on the history of fossil plants. Adolphe Brongniart concluded that the history of plants could roughly be divided into four parts. The first period was characterized by cryptogams. The second period was characterized by the appearance of the conifers. The third period brought emergence of the cycads, and the forth by the development of the flowering plants (such as the dicotyledons). The transitions between each of these periods was marked by sharp discontinuities in the fossil record, with gradual changes within clades of plants. Brongniart's work is the foundation of paleobotany and reinforced the theory that life on earth had a long and complex history, and different groups of plants and animals made their appearances in successive order. ^[20]

Catastrophism, uniformitarianism and the fossil record

In Cuvier's landmark 1796 paper on living and fossil elephants, he referred to a single catastrophe that destroyed life to be replaced by the current forms. As a result of his studies of extinct mammals, he realized that animals such as Palaeotherium had lived before the time of the Mammoths, which lead him to write in terms of multiple geological catastrophes that had wiped out a series of successive faunas.^[21] By 1830, a scientific consensus had formed around his ideas as a result of paleobotany and the dinosaur and marine reptile discoveries in Britain. ^[22] In Great Britain, where natural theology was very influential in the early 19th century, a group of geologists that included Buckland, and Robert Jameson insisted on explicitly linking the most recent of Cuvier's catastrophes to the biblical flood. Catastrophism had a religious overtone in Britain that was absent elsewhere.^[23]

Partly in response to what he saw as unsound and unscientific speculations by William Buckland and other practitioners of flood geology, Charles Lyell advocated the geological theory of uniformitarianism in his influential work Principles of Geology. ^[24] Lyell amassed evidence, both from his own field research and the work of others, that most geological features could be explained by the slow action of present day forces, such as vulcanism, earthquakes, erosion, and sedimentation rather than past catastrophic events. ^[25] Lyell also claimed that the apparent evidence for catastrophic changes in the fossil record, and even the appearance of directional succession in the history of life, were illusions caused by imperfections in that record. For instance he argued that the absence of birds and mammals from the earliest fossil strata was merely an imperfection in the fossil record attributable to the fact that marine organisms were more easily fossilized. ^[26] As evidence for these views Lyell pointed to the Stonesfield mammal as evidence that mammals had not necessarily been preceded by reptiles, and to the fact that certain Pleistocene strata showed a mixture of extinct and still surviving species, which he said showed that extinction occurred piecemeal rather than as a result of catastrophic events.^[27] Lyell was successful in convincing geologists of the idea that the geological features of the earth were largely due to the action of the same geologic forces that could be observed in the present day, acting over an extended period of time. He was not successful in gaining support for his view of the fossil record, which he believed did not support a theory of directional succession.^[28]

Transmutation of species and the fossil record

Jean Baptiste Lamarck used fossils in his arguments for his theory of the transmutation of species in the early 19th century.^[29] Fossil finds, and the emerging evidence that life had changed over time, fueled speculation on this topic during the next few decades.^[30] Robert Chambers used fossil evidence in his 1844 popular science book Vestiges of the Natural History of Creation, which advocated an evolutionary origin for the cosmos as well as for life on earth. Like Lamarck's theory it maintained that life had progressed from the simple to the complex^[31] These early evolutionary ideas were widely discussed in scientific circles but were not accepted into the scientific mainstream.^[32] Many of the critics of transmutational ideas used fossil evidence in their arguments. In the same paper that coined the term dinosaur Richard Owen pointed out that dinosaurs were at least as sophisticated and complex as modern reptiles, which he claimed contradicted transmutational theories.^[33] Hugh Miller would make a similar argument, pointing out that the fossil fish found in the Old Red Sandstone formation were fully as complex as any later fish, and not the primitive forms alleged by Vestiges.^[34] While these early evolutionary theories failed to become accepted as mainstream science, the debates over them would help pave the way for the acceptance of Darwin's theory of evoluton by natural selection a few years later.^[35]

Geological time scale and the history of life

Geologists such as Adam Sedgwick, and Roderick Murchison continued, despite disputes with other scientists, to make advances in stratigraphy. They described new geological epochs such as the Cambrian, the Silurian, the Devonian, and the Permian. By the early 1840s much of the geologic time scale had been developed. In 1841, John Phillips formally divided the geologic column into three major eras, the Paleozoic, Mesozoic, and Cenozoic, based on sharp breaks in the fossil record.^[36] He identified the three periods of the Mesozoic era and all the periods of the Paleozoic era except the Ordovician. His identification of the geological time scale is still used today.^[37] It remained a relative time scale with no method of assigning any of the periods' absolute dates. It was understood that not only had there been an "age of reptiles" preceding the current "age of mammals", but there had a time (during the Cambrian and the Silurian) when life had been restricted to the sea, and a time (prior to the Devonian) when invertebrates had been the largest and most complex forms of animal life.

Expansion and professionalization of geology and paleontology

This rapid progress in geology and paleontology during the 1830s and 1840s was aided by a growing international network of geologists and fossil specialists whose work was organized and reviewed by an increasing number of geological societies. Many of these geologists and paleontolgists were now paid professionals working for universities, museums and government geological surveys. The relatively high level of public support for the earth sciences was due to their cultural impact, and their proven economic value in helping to exploit mineral resources such as coal.^[38]

19th century after Darwin

Evolution

Charles Darwin's publication of the Origin of Species in 1859 was a watershed event in all the life sciences, especially paleontology. Fossils had played a role in the development of Darwin's theory. In particular he had been impressed by fossils he had collected in South America during the voyage of the Beagle of giant armadillos, giant sloths, and what at the time he thought were giant llamas that seemed to be related to species still living on the continent in modern times.^[39] The scientific debate that started immediately after the publication of Origin led to a concerted effort to look for transitional fossils and other evidence of evolution in the fossil record. There were two areas where early success attracted considerable public attention, the transition between reptiles and birds, and the evolution of the modern single-toed horse.^[40] In 1861 the first specimen of Archaeopteryx, an animal with both teeth and feathers and a mix of other reptilian and avian features, was discovered in a limestone quarry in Bavaria and described by Richard Owen. Another would be found in the late 1870s and put on display at a Museum in Berlin in 1881. Other primitive toothed birds were found by Othniel Marsh in Kansas in 1872. Marsh also discovered fossils of several primitive horses in the Western United States that helped trace the evolution of the horse from the small 5-toed Hyracotherium of the Eocene to the much larger single-toed modern horses of the genus Equus. Thomas Huxley would make extensive use of both the horse and bird fossils in his advocacy of evolution. Acceptance of evolution occurred rapidly in scientific circles, but acceptance of Darwin's proposed mechanism of natural selection as the driving force behind it was much less universal. In particular some paleontologists such as Edward Drinker Cope and Henry Fairfield Osborn preferred alternatives such as neo-Lamarckism, the inheritance of characteristics acquired during life, and orthogenesis, an innate drive to change in a particular direction, to explain what they perceived as linear trends in evolution.^[41]

There was also great interest in human evolution. Neanderthal fossils were discovered in 1856, but at the time it was not clear that they represented a different species from modern humans. Eugene Dubois created a sensation with his discovery of Java Man, the first fossil evidence of a species that seemed clearly intermediate between humans and apes, in 1891.

Developments in North America

A major development in the 2nd half of the 19th century was a rapid expansion of paleontology in North America. In 1858 Joseph Leidy described a Hadrosaurus skeleton, which was the first North American dinosaur to be described from good remains. However, it was the massive westward expansion of railroads, military bases, and settlements into Kansas and other parts of the Western United States following the American Civil War that really fueled the expansion of fossil collection.^[42] The result was an increased understanding of the natural history of north America, including the discovery of the Western Interior Sea that had covered Kansas and much of the rest of the Midwestern United States during parts of the Cretaceous, the discovery several important fossils of primitive birds and horses, and the discovery of a number of new dinosaur species including Allosaurus, Stegosaurus, and Triceratops. Much of this activity was part of a fierce personal and professional rivalry between two men, Othniel Marsh, and Edward Cope, which has become known as the Bone Wars.

Overview of developments in the 20th century

Developments in geology

Two 20th century developments in geology had a big effect on paleontology. The first was the development of radiometric dating, which allowed absolute dates to be assigned to the geologic timescale. The second was the theory of plate tectonics, which helped make sense of the geographical distribution of ancient life.

Geographical expansion of paleontology

During the 20th century paleontological exploration intensified everywhere and ceased to be a largely European and North American activity. In the 135 years between Buckland's first discovery and 1969 a total of 170 dinosaur genera were known. In the 25 years after 1969 that number increased to 315. Much of this increase was due to the examination of new rock exposures, particularly in previously little-explored areas in South America and Africa.^[43] Near the end of the century the opening of China to systematic exploration for fossils has yielded a wealth of material on dinosaurs and the origin of birds and mammals.^[44]

Mass extinctions

The 20th century saw a major renewal of interest in mass extinction events and their effect on the course of the history of life. This was particularly true after 1980 when Luis and Walter Alvarez put forward the Alvarez hypothesis claiming that an impact event caused the Cretaceous–Tertiary extinction event, which killed off the non-avian dinosaurs along with many other living things. Also in the early 1980's Jack Sepkoski and David M. Raup published papers with statistical analysis of the fossil record of marine invertebrates that revealed a pattern (possibly cyclical) of repeated mass extinctions with significant implications for the evolutionary history of life.

Evolutionary paths and theory

Throughout the 20th century new fossil finds continued to contribute to understanding the paths taken by evolution. Examples include major taxonomic transitions such as finds in Greenland, starting in the 1930’s (with more major finds in the 1980’s), of fossils illustrating the evolution of tetrapods from fish, and fossils in China during the 1990s that shed light on the dinosaur-bird relationship. Other events that have attracted considerable attention have included the discovery of a series of fossils in Pakistan that have shed light on whale evolution, and most famously of all a series of finds throughout the 20th century in Africa (starting with Taung child in 1924) and elsewhere have helped illuminate the course of human evolution. Increasingly, at the end of the century, the results of paleontology and molecular biology were being brought together to reveal detailed phylogenetic trees.

The results of paleontology have also contributed to the development of evolutionary theory. In 1944 George Gaylord Simpson published Tempo and Mode in Evolution, which used quantitative analysis to show that the fossil record was consistent with the branching, non-directional, patterns predicted by the advocates of evolution driven by natural selection and genetic drift rather than the linnear trends predicted by earlier advocates of neo-Lamarckism and orthogenesis. This integrated paleontology into the modern evolutionary synthesis.^[45] In 1972 Niles Eldredge and Stephen Jay Gould used fossil evidence to advocate the theory of punctuated equilibrium, which maintains that evolution is characterized by long periods of relative stasis and much shorter periods of relatively rapid change.

Cambrian explosion

One area of paleontology that has seen a lot of activity during the 1980’s, 1990’s and beyond is the study of the Cambrian explosion during which many of the various phyla of animals with their distinctive body plans first appear. The well-known Burgess Shale Cambrian fossil site was found in 1909 by Charles Doolittle Walcott, and another important site in Chengjiang China was found in 1912. However, new analysis in the 1980s by Harry B. Whittington, Derek Briggs, Simon Conway Morris and others sparked a renewed interest and a burst of activity including discovery of an important new fossil site, Sirius Passet, in Greenland, and the publication of a popular and controversial book, Wonderful Life by Stephen Jay Gould in 1989.

See also

• History of biology
• History of evolutionary thought
• History of geology
• History of science
• List of fossil sites (with link directory)

Notes

References

• Bowler, Peter J. (2003). Evolution:The History of an Idea. University of California Press. ISBN 0-52023693-9. 
• Desmond, Adrian (1975). "The Discovery of Marine Transgressions and the Explanation of Fossils in Antiquity". American Journal of Science, Volume 275.
• Larson, Edward J. (2004). Evolution: the remarkable history of scientific theory. Modern Library. ISBN 0-679-64288-9. 
• McGowan, Christopher (2001). The Dragon Seekers. Persus Publishing. ISBN 0-7382-0282-7. 
• Everhart, Michael J. (2005). Oceans of Kansas: A Natural History of the Western Interior Sea. Indiana University Press. ISBN 0-253-34547-2. 
• Needham, Joseph (1986). Science and Civilization in China: Volume 3, Mathematics and the Sciences of the Heavens and the Earth. Caves Books Ltd. ISBN 0-253-34547-2. 
• Robert Hooke (1665) em>Micrographia The Royal Society
• Rudwick, Martin J.S. (1997). Georges Cuvier, Fossil Bones, and Geological Catastrophes. The University of Chicago Press. ISBN 0-226-73106-5. 
• Rudwick, Martin J.S. (1972). The Meaning of Fossils. The University of Chicago Press. ISBN 0-226-73103-0. 

•  • [ e] History of biology
Fields and disciplines	Natural history • History of agriculture • History of medicine • History of anatomy • History of zoology (before Darwin)• History of plant systematics • History of geology • History of paleontology • History of evolutionary thought • History of zoology (after Darwin) • History of ecology • History of model organisms • History of phycology • History of genetics • History of biochemistry • History of agricultural science • History of molecular biology • History of molecular evolution • History of immunology • History of biotechnology
Institutions	Rothamsted Experimental Station • Pasteur Institute • Max Planck Society • Cold Spring Harbor Laboratory • Stazione Zoologica • Marine Biological Laboratory • Rockefeller University • Woods Hole Oceanographic Institute • Laboratory of Molecular Biology
Experiments	Griffith's experiment • Miller-Urey experiment • Luria-Delbrck experiment • Hershey-Chase experiment • Meselson-Stahl experiment • Crick, Brenner et al. experiment • Nirenberg and Matthaei experiment • Nirenberg and Leder experiment
Publications	On Generation and Corruption • Historia Plantarum • De humani corporis fabrica • De motu cordis • Micrographia • Systema Naturae • Philosophie Zoologique • Principles of Geology • Vestiges of Creation • The Origin of Species • "Experiments on Plant Hybridization" • The Descent of Man • "The Correlation Between Relatives on the Supposition of Mendelian Inheritance" • What is Life? • Genetics and the Origin of Species • "Molecular structure of Nucleic Acids"
Influential figures	Aristotle • Andreas Vesalius • William Harvey • Antonie van Leeuwenhoek • Carolus Linnaeus • Georges-Louis Leclerc, Comte de Buffon • Jean-Baptiste Lamarck • Alexander von Humboldt • Charles Lyell • Charles Darwin • Alfred Russel Wallace • Gregor Mendel • Louis Pasteur • Robert Koch • Ernst Haeckel • Ivan Pavlov • Hugo de Vries • E. B. Wilson • Thomas Hunt Morgan • Aleksandr Oparin • Alexander Fleming • J. B. S. Haldane • Sewall Wright • R. A. Fisher • Konrad Lorenz • Barbara McClintock • Theodosius Dobzhansky • Ernst Mayr • George Beadle • Seymour Benzer • Rosalind Franklin • James D. Watson • Francis Crick • Fred Sanger • Max Perutz • John Kendrew • Sydney Brenner • Joshua Lederberg • Walter Gilbert • Kary Mullis • Stephen Jay Gould • Lynn Margulis • Carl Woese • Jane Goodall
Related topics	History of science • History of medicine • Philosophy of biology • Timeline of biology and organic chemistry • Natural philosophy • Natural theology • Humboldtian science • Relationship between religion and science • Eugenics • Human Genome Project • Darwin Day • History of creationism • History of the creation-evolution controversy