Paleontology or
palaeontology (
//,
// or
//,
//) is the scientific study of life existent prior to, but sometimes including, the start of the
Holocene Epoch. It includes the study of fossils to determine organisms'
evolution and interactions with each other and their environments (their
paleoecology). Paleontological observations have been documented as far back as the 5th century BC. The science became established in the 18th century as a result of
Georges Cuvier's work on
comparative anatomy, and developed rapidly in the 19th century. The term itself originates from
Greek παλαιός,
palaios, i.e. "old, ancient", ὄν,
on (
gen. ontos), i.e. "being, creature" and λόγος,
logos, i.e. "speech, thought, study".
[1]
Paleontology lies on the border between
biology and
geology, but differs from
archaeology in that it excludes the study of morphologically modern humans. It now uses techniques drawn from a wide range of sciences, including
biochemistry,
mathematics and
engineering. Use of all these techniques has enabled paleontologists to discover much of the
evolutionary history of life, almost all the way back to when
Earth became capable of supporting life, about
3,800 million years ago. As knowledge has increased, paleontology has developed specialized sub-divisions, some of which focus on different types of
fossil organisms while others study
ecology and environmental history, such as
ancient climates.
Body fossils and
trace fossils are the principal types of evidence about ancient life, and
geochemical evidence has helped to decipher the evolution of life before there were organisms large enough to leave body fossils. Estimating the dates of these remains is essential but difficult: sometimes adjacent rock layers allow
radiometric dating, which provides
absolute dates that are accurate to within 0.5%, but more often paleontologists have to rely on relative dating by solving the "
jigsaw puzzles" of
biostratigraphy. Classifying ancient organisms is also difficult, as many do not fit well into the
Linnean taxonomy that is commonly used for classifying living organisms, and paleontologists more often use
cladistics to draw up evolutionary "family trees". The final quarter of the 20th century saw the development of
molecular phylogenetics, which investigates how closely organisms are related by measuring how similar the
DNA is in their
genomes. Molecular phylogenetics has also been used to estimate the dates when species diverged, but there is controversy about the reliability of the
molecular clock on which such estimates depend.
§Overview
The simplest definition is "the study of ancient life".
[2] Paleontology seeks information about several aspects of past organisms: "their identity and origin, their environment and evolution, and what they can tell us about the Earth's organic and inorganic past".
[3]
§A historical science
Paleontology is one of the historical sciences, along with
archaeology,
geology,
astronomy,
cosmology,
philology and
history itself.
[4] This means that it aims to describe phenomena of the past and reconstruct their causes.
[5] Hence it has three main elements: description of the phenomena; developing a general theory about the causes of various types of change; and applying those theories to specific facts.
[4]
When trying to explain past phenomena, paleontologists and other historical scientists often construct a set of hypotheses about the causes and then look for a
smoking gun, a piece of evidence that indicates that one hypotheses is a better explanation than others. Sometimes the smoking gun is discovered by a fortunate accident during other research. For example, the discovery by
Luis Alvarez and
Walter Alvarez of an
iridium-rich layer at the
Cretaceous–
Tertiary boundary made
asteroid impact and
volcanism the most favored explanations for the
Cretaceous–Paleogene extinction event.
[5]
The other main type of science is experimental science, which is often said to work by conducting
experiments to
disprove hypotheses about the workings and causes of natural phenomena – note that this approach cannot confirm a hypothesis is correct, since some later experiment may disprove it. However, when confronted with totally unexpected phenomena, such as the first evidence for invisible
radiation, experimental scientists often use the same approach as historical scientists: construct a set of hypotheses about the causes and then look for a "smoking gun".
[5]
§Related sciences
Paleontology lies on the boundary between
biology and
geology since paleontology focuses on the record of past life but its main source of evidence is
fossils, which are found in rocks.
[6] For historical reasons paleontology is part of the geology departments of many universities, because in the 19th century and early 20th century geology departments found paleontological evidence important for estimating the ages of rocks while biology departments showed little interest.
[7]
Paleontology also has some overlap with
archaeology, which primarily works with objects made by humans and with human remains, while paleontologists are interested in the characteristics and evolution of humans as organisms. When dealing with evidence about humans, archaeologists and paleontologists may work together – for example paleontologists might identify animal or plant fossils around an
archaeological site, to discover what the people who lived there ate; or they might analyze the climate at the time when the site was inhabited by humans.
[8]
Analyses using
engineering techniques show that
Tyrannosaurus had a devastating bite, but raise doubts about how fast it could move.
In addition paleontology often uses techniques derived from other sciences, including biology,
osteology,
ecology,
chemistry,
physics and
mathematics.
[2] For example
geochemical signatures from rocks may help to discover when life first arose on Earth,
[9] and analyses of
carbon isotope ratios may help to identify climate changes and even to explain major transitions such as the
Permian–Triassic extinction event.
[10] A relatively recent discipline,
molecular phylogenetics, often helps by using comparisons of different modern organisms'
DNA and
RNA to re-construct evolutionary "family trees"; it has also been used to estimate the dates of important evolutionary developments, although this approach is controversial because of doubts about the reliability of the "
molecular clock".
[11] Techniques developed in
engineering have been used to analyse how ancient organisms might have worked, for example how fast
Tyrannosaurus could move and how powerful its bite was.
[12][13]
A combination of paleontology, biology, and archaeology,
paleoneurology is the study of endocranial casts (or endocasts) of species related to humans to learn about the evolution of human brains.
[14]
Paleontology even contributes to
astrobiology, the investigation of possible life on other
planets, by developing models of how life may have arisen and by providing techniques for detecting evidence of life.
[15]
§Subdivisions
As knowledge has increased, paleontology has developed specialised subdivisions.
[16] Vertebrate paleontology concentrates on
fossils of
vertebrates, from the earliest
fish to the immediate ancestors of modern
mammals.
Invertebrate paleontology deals with fossils of
invertebrates such as
molluscs,
arthropods,
annelid worms and
echinoderms.
Paleobotany focuses on the study of fossil
plants, but traditionally includes the study of fossil
algae and
fungi.
Palynology, the study of
pollen and
spores produced by land plants and
protists, straddles the border between paleontology and
botany, as it deals with both living and fossil organisms.
Micropaleontology deals with all microscopic fossil organisms, regardless of the group to which they belong.
[17]
Instead of focusing on individual organisms,
paleoecology examines the interactions between different organisms, such as their places in
food chains, and the two-way interaction between organisms and their environment.
[18] One example is the development of
oxygenic photosynthesis by
bacteria, which hugely increased the productivity and diversity of
ecosystems.
[19] This also caused the
oxygenation of the atmosphere. Together, these were a prerequisite for the evolution of the most complex
eucaryotic cells, from which all
multicellular organisms are built.
[20]
Paleoclimatology, although sometimes treated as part of paleoecology,
[17] focuses more on the history of Earth's climate and the mechanisms that have changed it
[21] – which have sometimes included
evolutionary developments, for example the rapid expansion of land plants in the
Devonian period removed more
carbon dioxide from the atmosphere, reducing the
greenhouse effect and thus helping to cause an
ice age in the
Carboniferous period.
[22]
Biostratigraphy, the use of fossils to work out the chronological order in which rocks were formed, is useful to both paleontologists and geologists.
[23] Biogeography studies the spatial distribution of organisms, and is also linked to geology, which explains how Earth's geography has changed over time.
[24]
§Sources of evidence
§Body fossils
Fossils of organisms' bodies are usually the most informative type of evidence. The most common types are wood, bones, and shells.
[25] Fossilisation is a rare event, and most fossils are destroyed by
erosion or
metamorphism before they can be observed. Hence the fossil record is very incomplete, increasingly so further back in time. Despite this, it is often adequate to illustrate the broader patterns of life's history.
[26] There are also biases in the fossil record: different environments are more favorable to the preservation of different types of organism or parts of organisms.
[27] Further, only the parts of organisms that were already
mineralised are usually preserved, such as the shells of molluscs. Since most animal species are soft-bodied, they decay before they can become fossilised. As a result, although there are 30-plus
phyla of living animals, two-thirds have never been found as fossils.
[28]
Occasionally, unusual environments may preserve soft tissues. These
lagerstätten allow paleontologists to examine the internal anatomy of animals that in other sediments are represented only by shells, spines, claws, etc. – if they are preserved at all. However, even lagerstätten present an incomplete picture of life at the time. The majority of organisms living at the time are probably not represented because lagerstätten are restricted to a narrow range of environments, e.g. where soft-bodied organisms can be preserved very quickly by events such as mudslides; and the exceptional events that cause quick burial make it difficult to study the normal environments of the animals.
[29] The sparseness of the fossil record means that organisms are expected to exist long before and after they are found in the fossil record – this is known as the
Signor-Lipps effect.
[30]
§Trace fossils
Trace fossils consist mainly of tracks and burrows, but also include
coprolites (fossil
feces) and marks left by feeding.
[25][31] Trace fossils are particularly significant because they represent a data source that is not limited to animals with easily fossilized hard parts, and they reflect organisms' behaviours. Also many traces date from significantly earlier than the body fossils of animals that are thought to have been capable of making them.
[32]
Whilst exact assignment of trace fossils to their makers is generally impossible, traces may for example provide the earliest physical evidence of the appearance of moderately complex animals (comparable to
earthworms).
[31]
§Geochemical observations
Geochemical observations may help to deduce the global level of biological activity, or the affinity of a certain fossil. For example geochemical features of rocks may reveal when life first arose on Earth,
[9] and may provide evidence of the presence of
eucaryotic cells, the type from which all
multicellular organisms are built.
[33] Analyses of
carbon isotope ratios may help to explain major transitions such as the
Permian–Triassic extinction event.
[10]
§Classifying ancient organisms
Naming groups of organisms in a way that is clear and widely agreed is important, as some disputes in paleontology have been based just on misunderstandings over names.
[35] Linnean taxonomy is commonly used for classifying living organisms, but runs into difficulties when dealing with newly discovered organisms that are significantly different from known ones. For example: it is hard to decide at what level to place a new higher-level grouping, e.g.
genus or
family or
order; this is important since the Linnean rules for naming groups are tied to their levels, and hence if a group is moved to a different level it must be renamed.
[36]
Paleontologists generally use approaches based on
cladistics, a technique for working out the evolutionary "family tree" of a set of organisms.
[35] It works by the logic that, if groups B and C have more similarities to each other than either has to group A, then B and C are more closely related to each other than either is to A. Characters that are compared may be
anatomical, such as the presence of a
notochord, or
molecular, by comparing sequences of
DNA or
proteins. The result of a successful analysis is a hierarchy of clades – groups that share a common ancestor. Ideally the "family tree" has only two branches leading from each node ("junction"), but sometimes there is too little information to achieve this and paleontologists have to make do with junctions that have several branches. The cladistic technique is sometimes fallible, as some features, such as wings or
camera eyes, evolved more than once,
convergently – this must be taken into account in analyses.
[34]
Evolutionary developmental biology, commonly abbreviated to "Evo Devo", also helps paleontologists to produce "family trees". For example the
embryological development of some modern
brachiopods suggests that brachiopods may be descendants of the
halkieriids, which became extinct in the
Cambrian period.
[37]
§Estimating the dates of organisms
Paleontology seeks to map out how living things have changed through time. A substantial hurdle to this aim is the difficulty of working out how old fossils are. Beds that preserve fossils typically lack the radioactive elements needed for
radiometric dating. This technique is our only means of giving rocks greater than about 50 million years old an absolute age, and can be accurate to within 0.5% or better.
[38] Although radiometric dating requires very careful laboratory work, its basic principle is simple: the rates at which various radioactive elements
decay are known, and so the ratio of the radioactive element to the element into which it decays shows how long ago the radioactive element was incorporated into the rock. Radioactive elements are common only in rocks with a volcanic origin, and so the only fossil-bearing rocks that can be dated radiometrically are a few volcanic ash layers.
[38]
Consequently, paleontologists must usually rely on
stratigraphy to date fossils. Stratigraphy is the science of deciphering the "layer-cake" that is the
sedimentary record, and has been compared to a
jigsaw puzzle.
[39] Rocks normally form relatively horizontal layers, with each layer younger than the one underneath it. If a fossil is found between two layers whose ages are known, the fossil's age must lie between the two known ages.
[40] Because rock sequences are not continuous, but may be broken up by
faults or periods of
erosion, it is very difficult to match up rock beds that are not directly next to one another. However, fossils of species that survived for a relatively short time can be used to link up isolated rocks: this technique is called
biostratigraphy. For instance, the conodont
Eoplacognathus pseudoplanus has a short range in the Middle Ordovician period.
[41] If rocks of unknown age are found to have traces of
E. pseudoplanus, they must have a mid-Ordovician age. Such
index fossils must be distinctive, be globally distributed and have a short time range to be useful. However, misleading results are produced if the index fossils turn out to have longer fossil ranges than first thought.
[42] Stratigraphy and biostratigraphy can in general provide only relative dating (
A was before
B), which is often sufficient for studying evolution. However, this is difficult for some time periods, because of the problems involved in matching up rocks of the same age across different
continents.
[43]
Family-tree relationships may also help to narrow down the date when lineages first appeared. For instance, if fossils of B or C date to X million years ago and the calculated "family tree" says A was an ancestor of B and C, then A must have evolved more than X million years ago.
It is also possible to estimate how long ago two living clades diverged – i.e. approximately how long ago their last common ancestor must have lived – by assuming that DNA
mutations accumulate at a constant rate. These "
molecular clocks", however, are fallible, and provide only a very approximate timing: for example, they are not sufficiently precise and reliable for estimating when the groups that feature in the
Cambrian explosion first evolved,
[44] and estimates produced by different techniques may vary by a factor of two.
[11]
§Overview of the history of life
The evolutionary history of life stretches back to over
3,000 million years ago, possibly as far as
3,800 million years ago.
Earth formed about
4,570 million years ago and, after a collision that formed the
Moon about 40 million years later, may have cooled quickly enough to have oceans and an atmosphere about
4,440 million years ago.
[45] However there is evidence on the Moon of a
Late Heavy Bombardment from
4,000 to 3,800 million years ago. If, as seem likely, such a bombardment struck Earth at the same time, the first atmosphere and oceans may have been stripped away.
[46] The oldest clear evidence of life on Earth dates to
3,000 million years ago, although there have been reports, often disputed, of
fossil bacteria from
3,400 million years ago and of geochemical evidence for the presence of life
3,800 million years ago.
[9][47] Some scientists have proposed that life on Earth was
"seeded" from elsewhere,
[48] but most research concentrates on various explanations of how life could have
arisen independently on Earth.
[49]
For about 2,000 million years
microbial mats, multi-layered colonies of different types of bacteria, were the dominant life on Earth.
[51] The evolution of
oxygenic photosynthesis enabled them to play the major role in the
oxygenation of the atmosphere[52] from about
2,400 million years ago. This change in the atmosphere increased their effectiveness as nurseries of evolution.
[53] While
eukaryotes, cells with complex internal structures, may have been present earlier, their evolution speeded up when they acquired the ability to transform oxygen from a
poison to a powerful source of energy in their
metabolism. This innovation may have come from primitive eukaryotes capturing oxygen-powered bacteria as
endosymbionts and transforming them into
organelles called
mitochondria.
[54] The earliest evidence of complex eukaryotes with organelles such as mitochondria, dates from
1,850 million years ago.
[20]
Multicellular life is composed only of eukaryotic cells, and the earliest evidence for it is the
Francevillian Group Fossils from
2,100 million years ago,
[55] although specialization of cells for different functions first appears between
1,430 million years ago (a possible
fungus) and
1,200 million years ago (a probable
red alga).
Sexual reproduction may be a prerequisite for specialization of cells, as an asexual multicellular organism might be at risk of being taken over by rogue cells that retain the ability to reproduce.
[56][57]
The earliest known
animals are
cnidarians from about
580 million years ago, but these are so modern-looking that the earliest animals must have appeared before then.
[58] Early fossils of animals are rare because they did not develop
mineralized hard parts that fossilize easily until about
548 million years ago.
[59] The earliest modern-looking
bilaterian animals appear in the Early
Cambrian, along with several "weird wonders" that bear little obvious resemblance to any modern animals. There is a long-running debate about whether this
Cambrian explosion was truly a very rapid period of evolutionary experimentation; alternative views are that modern-looking animals began evolving earlier but fossils of their precursors have not yet been found, or that the "weird wonders" are
evolutionary "aunts" and "cousins" of modern groups.
[60] Vertebrates remained an obscure group until the first fish with jaws appeared in the Late
Ordovician.
[61][62]
The spread of life from water to land required organisms to solve several problems, including protection against drying out and supporting themselves against
gravity.
[63][64][65] The earliest evidence of land plants and land invertebrates date back to about
476 million years ago and
490 million years ago respectively.
[64][66] The lineage that produced land vertebrates evolved later but very rapidly between
370 million years ago and
360 million years ago;
[67] recent discoveries have overturned earlier ideas about the history and driving forces behind their evolution.
[68] Land plants were so successful that they caused an
ecological crisis in the Late
Devonian, until the evolution and spread of fungi that could digest dead wood.
[22]
At about 13 centimetres (5.1 in) the Early Cretaceous
Yanoconodon was longer than the average mammal of the time.
[69]
During the
Permian period
synapsids, including the ancestors of
mammals, may have dominated land environments,
[71] but the
Permian–Triassic extinction event 251 million years ago came very close to wiping out complex life.
[72] The extinctions were apparently fairly sudden, at least among vertebrates.
[73] During the slow recovery from this catastrophe a previously obscure group,
archosaurs, became the most abundant and diverse terrestrial vertebrates. One archosaur group, the
dinosaurs, were the dominant land vertebrates for the rest of the
Mesozoic,
[74] and
birds evolved from one group of dinosaurs.
[70] During this time mammals' ancestors survived only as small, mainly nocturnal
insectivores, but this apparent set-back may have accelerated the development of mammalian traits such as
endothermy and
hair.
[75] After the
Cretaceous–Paleogene extinction event 65 million years ago killed off the non-avian dinosaurs – birds are the only surviving dinosaurs – mammals increased rapidly in size and diversity, and some took to the air and the sea.
[76][77][78]
Fossil evidence indicates that
flowering plants appeared and rapidly diversified in the Early
Cretaceous, between
130 million years ago and
90 million years ago.
[79] Their rapid rise to dominance of terrestrial ecosystems is thought to have been propelled by
coevolution with
pollinating insects.
[80] Social insects appeared around the same time and, although they account for only small parts of the insect "family tree", now form over 50% of the total mass of all insects.
[81]
Humans evolved from a lineage of upright-walking
apes whose earliest fossils date from over
6 million years ago.
[82] Although early members of this lineage had
chimp-sized
brains, about 25% as big as modern humans', there are signs of a steady increase in brain size after about
3 million years ago.
[83] There is a long-running debate about whether
modern humans are descendants of a
single small population in Africa, which then migrated all over the world less than 200,000 years ago and replaced previous
hominine species, or
arose worldwide at the same time as a result of
interbreeding.
[84]
§Mass extinctions
Life on earth has suffered occasional mass extinctions at least since
542 million years ago. Although they are disasters at the time, mass extinctions have sometimes accelerated the evolution of
life on earth. When dominance of particular
ecological niches passes from one group of organisms to another, it is rarely because the new dominant group is "superior" to the old and usually because an extinction event eliminates the old dominant group and makes way for the new one.
[85][86]
The fossil record appears to show that the rate of extinction is slowing down, with both the gaps between mass extinctions becoming longer and the average and background rates of extinction decreasing. However, it is not certain whether the actual rate of extinction has altered, since both of these observations could be explained in several ways:
[87]
- The oceans may have become more hospitable to life over the last 500 million years and less vulnerable to mass extinctions: dissolved oxygen became more widespread and penetrated to greater depths; the development of life on land reduced the run-off of nutrients and hence the risk of eutrophication and anoxic events; marine ecosystems became more diversified so that food chains were less likely to be disrupted.[88][89]
- Reasonably complete fossils are very rare, most extinct organisms are represented only by partial fossils, and complete fossils are rarest in the oldest rocks. So paleontologists have mistakenly assigned parts of the same organism to different genera, which were often defined solely to accommodate these finds – the story of Anomalocaris is an example of this.[90] The risk of this mistake is higher for older fossils because these are often unlike parts of any living organism. Many "superfluous" genera are represented by fragments that are not found again, and these "superfluous" genera appear to become extinct very quickly.[87]
"Well-defined" genera
Trend line
Other mass extinctions
Million years ago
Thousands of genera
Phanerozoic biodiversity as shown by the fossil record
Biodiversity in the fossil record, which is
-
- "the number of distinct genera alive at any given time; that is, those whose first occurrence predates and whose last occurrence postdates that time"[91]
shows a different trend: a fairly swift rise from
542 to 400 million years ago, a slight decline from
400 to 200 million years ago, in which the devastating
Permian–Triassic extinction event is an important factor, and a swift rise from
200 million years ago to the present.
[91]
§History of paleontology
Although paleontology became established around 1800, earlier thinkers had noticed aspects of the
fossil record. The ancient Greek
philosopher Xenophanes (570–480 BC) concluded from fossil sea shells that some areas of land were once under water.
[92] During the
Middle Ages the Persian naturalist
Ibn Sina, known as
Avicenna in Europe, discussed fossils and proposed a theory of petrifying fluids on which
Albert of Saxony elaborated in the 14th century.
[93] The Chinese naturalist
Shen Kuo (1031–1095) proposed a theory of climate change based on the presence of
petrified bamboo in regions that in his time were too dry for bamboo.
[94]
In
early modern Europe, the systematic study of fossils emerged as an integral part of the changes in
natural philosophy that occurred during the
Age of Reason. At the end of the 18th century
Georges Cuvier's work established
comparative anatomy as a scientific discipline and, by proving that some fossil animals resembled no living ones, demonstrated that animals could become
extinct, leading to the emergence of paleontology.
[95] The expanding knowledge of the fossil record also played an increasing role in the development of
geology, particularly
stratigraphy.
[96]
The first half of the 19th century saw geological and paleontological activity become increasingly well organized with the growth of geologic societies and museums
[97][98] and an increasing number of professional geologists and fossil specialists. Interest increased for reasons that were not purely scientific, as geology and paleontology helped industrialists to find and exploit natural resources such as
coal.
[99]
This contributed to a rapid increase in knowledge about the history of life on Earth and to progress in the definition of the
geologic time scale, largely based on fossil evidence. In 1822 Henri Marie Ducrotay de Blanville, editor of
Journal de Phisique, coined the word "palaeontology" to refer to the study of ancient living organisms through fossils.
[100] As knowledge of life's history continued to improve, it became increasingly obvious that there had been some kind of successive order to the development of life. This encouraged early evolutionary theories on the
transmutation of species.
[101] 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.
[101]
The last half of the 19th century saw a tremendous expansion in paleontological activity, especially in
North America.
[103] The trend continued in the 20th century with additional regions of the Earth being opened to systematic fossil collection. Fossils found in China near the end of the 20th century have been particularly important as they have provided new information about the earliest evolution of
animals, early
fish,
dinosaurs and the evolution of birds.
[104] The last few decades of the 20th century saw a renewed interest in
mass extinctions and their role in the evolution of life on Earth.
[105] There was also a renewed interest in the
Cambrian explosion that apparently saw the development of the body plans of most animal
phyla. The discovery of fossils of the
Ediacaran biota and developments in
paleobiology extended knowledge about the history of life back far before the Cambrian.
[60]
Increasing awareness of
Gregor Mendel's pioneering work in
genetics led first to the development of
population genetics and then in the mid-20th century to the
modern evolutionary synthesis, which explains
evolution as the outcome of events such as
mutations and
horizontal gene transfer, which provide
genetic variation, with
genetic drift and
natural selection driving changes in this variation over time.
[106] Within the next few years the role and operation of
DNA in genetic inheritance were discovered, leading to what is now known as the
"Central Dogma" of molecular biology.
[107] In the 1960s
molecular phylogenetics, the investigation of evolutionary "family trees" by techniques derived from
biochemistry, began to make an impact, particularly when it was proposed that the human lineage had diverged from
apes much more recently than was generally thought at the time.
[108] Although this early study compared
proteins from apes and humans, most molecular phylogenetics research is now based on comparisons of
RNA and
DNA.
[109]
A study of embryos by Leonardo da Vinci
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