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Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the
Holocene The Holocene ( ) is the current geological epoch. It began approximately 11,650 cal years Before Present (), after the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene togeth ...
epoch In chronology and periodization, an epoch or reference epoch is an instant in time chosen as the origin of a particular calendar era. The "epoch" serves as a reference point from which time is measured. The moment of epoch is usually decided ...
(roughly 11,700 years before present). It includes the study of
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s to classify
organism In biology, an organism () is any living system that functions as an individual entity. All organisms are composed of cells ( cell theory). Organisms are classified by taxonomy into groups such as multicellular animals, plants, and fu ...
s and study their
interactions Interaction is action that occurs between two or more objects, with broad use in philosophy and the sciences. It may refer to: Science * Interaction hypothesis, a theory of second language acquisition * Interaction (statistics) * 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 Jean Léopold Nicolas Frédéric, Baron Cuvier (; 23 August 1769 – 13 May 1832), known as Georges Cuvier, was a French naturalist and zoologist, sometimes referred to as the "founding father of paleontology". Cuvier was a major figure in na ...
's work on
comparative anatomy Comparative anatomy is the study of similarities and differences in the anatomy of different species. It is closely related to evolutionary biology and phylogeny (the evolution of species). The science began in the classical era, continuing in ...
, and developed rapidly in the 19th century. The term itself originates from Greek (, "old, ancient"), (, (
gen. The Book of Genesis (from Greek ; Hebrew: בְּרֵאשִׁית ''Bəreʾšīt'', "In hebeginning") is the first book of the Hebrew Bible and the Christian Old Testament. Its Hebrew name is the same as its first word, ( "In the beginning"). ...
), "being, creature"), and (, "speech, thought, study"). Paleontology lies on the border between
biology Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells that process hereditary ...
and
geology Geology () is a branch of natural science concerned with Earth and other Astronomical object, astronomical objects, the features or rock (geology), rocks of which it is composed, and the processes by which they change over time. Modern geology ...
, but differs from
archaeology Archaeology or archeology is the scientific study of human activity through the recovery and analysis of material culture. The archaeological record consists of artifacts, architecture, biofacts or ecofacts, sites, and cultural landsc ...
in that it excludes the study of anatomically modern humans. It now uses techniques drawn from a wide range of sciences, including
biochemistry Biochemistry or biological chemistry is the study of chemical processes within and relating to living organisms. A sub-discipline of both chemistry and biology, biochemistry may be divided into three fields: structural biology, enzymology and ...
,
mathematic Mathematics is an area of knowledge that includes the topics of numbers, formulas and related structures, shapes and the spaces in which they are contained, and quantities and their changes. These topics are represented in modern mathematics ...
s, 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, nearly 4 billion years ago. As knowledge has increased, paleontology has developed specialised sub-divisions, some of which focus on different types of fossil organisms while others study
ecology Ecology () is the study of the relationships between living organisms, including humans, and their physical environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere level. Ecology overl ...
and environmental history, such as ancient climates. Body fossils and trace fossils are the principal types of evidence about ancient life, and
geochemical Geochemistry is the science that uses the tools and principles of chemistry to explain the mechanisms behind major geological systems such as the Earth's crust and its oceans. The realm of geochemistry extends beyond the Earth, encompassing the e ...
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 Radiometric dating, radioactive dating or radioisotope dating is a technique which is used to date materials such as rocks or carbon, in which trace radioactive impurities were selectively incorporated when they were formed. The method compares ...
, 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 puzzle A jigsaw puzzle is a tiling puzzle that requires the assembly of often irregularly shaped interlocking and mosaiced pieces, each of which typically has a portion of a picture. When assembled, the puzzle pieces produce a complete picture. In t ...
s" of
biostratigraphy Biostratigraphy is the branch of stratigraphy which focuses on correlating and assigning relative ages of rock strata by using the fossil assemblages contained within them.Hine, Robert. “Biostratigraphy.” ''Oxford Reference: Dictionary of ...
(arrangement of rock layers from youngest to oldest). Classifying ancient organisms is also difficult, as many do not fit well into the
Linnaean taxonomy Linnaean taxonomy can mean either of two related concepts: # The particular form of biological classification (taxonomy) set up by Carl Linnaeus, as set forth in his ''Systema Naturae'' (1735) and subsequent works. In the taxonomy of Linnaeus t ...
classifying living organisms, and paleontologists more often use
cladistics Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups (" clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived cha ...
to draw up evolutionary "family trees". The final quarter of the 20th century saw the development of
molecular phylogenetics Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
, which investigates how closely organisms are related by measuring the similarity of the DNA in their
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ...
s. Molecular phylogenetics has also been used to estimate the dates when species diverged, but there is controversy about the reliability of the
molecular clock The molecular clock is a figurative term for a technique that uses the mutation rate of biomolecules to deduce the time in prehistory when two or more life forms diverged. The biomolecular data used for such calculations are usually nucleo ...
on which such estimates depend.


Overview

The simplest definition of "paleontology" is "the study of ancient life". The field 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".


Historical science

William Whewell William Whewell ( ; 24 May 17946 March 1866) was an English polymath, scientist, Anglican priest, philosopher, theologian, and historian of science. He was Master of Trinity College, Cambridge. In his time as a student there, he achieved ...
(1794–1866) classified paleontology as one of the historical sciences, along with
archaeology Archaeology or archeology is the scientific study of human activity through the recovery and analysis of material culture. The archaeological record consists of artifacts, architecture, biofacts or ecofacts, sites, and cultural landsc ...
, geology,
astronomy Astronomy () is a natural science that studies celestial objects and phenomena. It uses mathematics, physics, and chemistry in order to explain their origin and evolution. Objects of interest include planets, moons, stars, nebulae, g ...
,
cosmology Cosmology () is a branch of physics and metaphysics dealing with the nature of the universe. The term ''cosmology'' was first used in English in 1656 in Thomas Blount's ''Glossographia'', and in 1731 taken up in Latin by German philosopher ...
,
philology Philology () is the study of language in oral and written historical sources; it is the intersection of textual criticism, literary criticism, history, and linguistics (with especially strong ties to etymology). Philology is also defined as ...
and history itself: paleontology aims to describe phenomena of the past and to reconstruct their causes. Hence it has three main elements: description of past phenomena; developing a general theory about the causes of various types of change; and applying those theories to specific facts. When trying to explain the past, paleontologists and other historical scientists often construct a set of one or more
hypotheses A hypothesis (plural hypotheses) is a proposed explanation for a phenomenon. For a hypothesis to be a scientific hypothesis, the scientific method requires that one can test it. Scientists generally base scientific hypotheses on previous obse ...
about the causes and then look for a "
smoking gun The term "smoking gun" is a reference to an object or fact that serves as conclusive evidence of a crime or similar act, just short of being caught '' in flagrante delicto''. "Smoking gun" refers to the strongest kind of circumstantial evidenc ...
", a piece of evidence that strongly accords with one hypothesis over any others. Sometimes researchers discover a "smoking gun" by a fortunate accident during other research. For example, the 1980 discovery by Luis and
Walter Alvarez Walter Alvarez (born October 3, 1940) is a professor in the Earth and Planetary Science department at the University of California, Berkeley. He is most widely known for the theory that dinosaurs were killed by an asteroid impact, developed in ...
of
iridium Iridium is a chemical element with the symbol Ir and atomic number 77. A very hard, brittle, silvery-white transition metal of the platinum group, it is considered the second-densest naturally occurring metal (after osmium) with a density o ...
, a mainly extraterrestrial metal, in the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
Tertiary Tertiary ( ) is a widely used but obsolete term for the geologic period from 66 million to 2.6 million years ago. The period began with the demise of the non-avian dinosaurs in the Cretaceous–Paleogene extinction event, at the start ...
boundary layer made
asteroid impact An impact event is a collision between astronomical objects causing measurable effects. Impact events have physical consequences and have been found to regularly occur in planetary systems, though the most frequent involve asteroids, comets or me ...
the most favored explanation for the
Cretaceous–Paleogene extinction event The Cretaceous–Paleogene (K–Pg) extinction event (also known as the Cretaceous–Tertiary extinction) was a sudden mass extinction of three-quarters of the plant and animal species on Earth, approximately 66 million years ago. With the ...
– although debate continues about the contribution of volcanism. A complementary approach to developing scientific knowledge,
experimental science An experiment is a procedure carried out to support or refute a hypothesis, or determine the efficacy or likelihood of something previously untried. Experiments provide insight into cause-and-effect by demonstrating what outcome occurs when ...
, is often said to work by conducting experiments to ''disprove'' hypotheses about the workings and causes of natural phenomena. This approach cannot prove a hypothesis, since some later experiment may disprove it, but the accumulation of failures to disprove is often compelling evidence in favor. However, when confronted with totally unexpected phenomena, such as the first evidence for invisible
radiation In physics, radiation is the emission or transmission of energy in the form of waves or particles through space or through a material medium. This includes: * ''electromagnetic radiation'', such as radio waves, microwaves, infrared, visi ...
, 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".


Related sciences

Paleontology lies between
biology Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells that process hereditary ...
and geology since it focuses on the record of past life, but its main source of evidence is
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s in rocks. For historical reasons, paleontology is part of the geology department at many universities: in the 19th and early 20th centuries, geology departments found fossil evidence important for dating rocks, while biology departments showed little interest. Paleontology also has some overlap with
archaeology Archaeology or archeology is the scientific study of human activity through the recovery and analysis of material culture. The archaeological record consists of artifacts, architecture, biofacts or ecofacts, sites, and cultural landsc ...
, which primarily works with objects made by humans and with human remains, while paleontologists are interested in the characteristics and evolution of humans as a species. 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 An archaeological site is a place (or group of physical sites) in which evidence of past activity is preserved (either prehistoric or historic or contemporary), and which has been, or may be, investigated using the discipline of archaeology an ...
, to discover the people who lived there, and what they ate; or they might analyze the climate at the time of habitation. In addition, paleontology often borrows techniques from other sciences, including biology,
osteology Osteology () is the scientific study of bones, practised by osteologists. A subdiscipline of anatomy, anthropology, and paleontology, osteology is the detailed study of the structure of bones, skeletal elements, teeth, microbone morphology, func ...
, ecology,
chemistry Chemistry is the scientific study of the properties and behavior of matter. It is a natural science that covers the elements that make up matter to the compounds made of atoms, molecules and ions: their composition, structure, proper ...
,
physics Physics is the natural science that studies matter, its fundamental constituents, its motion and behavior through space and time, and the related entities of energy and force. "Physical science is that department of knowledge which ...
and mathematics. For example,
geochemical Geochemistry is the science that uses the tools and principles of chemistry to explain the mechanisms behind major geological systems such as the Earth's crust and its oceans. The realm of geochemistry extends beyond the Earth, encompassing the e ...
signatures from rocks may help to discover when life first arose on Earth, and analyses of
carbon Carbon () is a chemical element with the symbol C and atomic number 6. It is nonmetallic and tetravalent—its atom making four electrons available to form covalent chemical bonds. It belongs to group 14 of the periodic table. Carbon ma ...
isotope ratios may help to identify climate changes and even to explain major transitions such as the
Permian–Triassic extinction event The Permian–Triassic (P–T, P–Tr) extinction event, also known as the Latest Permian extinction event, the End-Permian Extinction and colloquially as the Great Dying, formed the boundary between the Permian and Triassic geologic periods, ...
. A relatively recent discipline,
molecular phylogenetics Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
, compares the DNA and RNA of modern organisms to re-construct the "family trees" of their evolutionary ancestors. 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 The molecular clock is a figurative term for a technique that uses the mutation rate of biomolecules to deduce the time in prehistory when two or more life forms diverged. The biomolecular data used for such calculations are usually nucleo ...
". Techniques from engineering have been used to analyse how the bodies of ancient organisms might have worked, for example the running speed and bite strength of ''
Tyrannosaurus ''Tyrannosaurus'' is a genus of large theropoda, theropod dinosaur. The species ''Tyrannosaurus rex'' (''rex'' meaning "king" in Latin), often called ''T. rex'' or colloquially ''T-Rex'', is one of the best represented theropods. ''Tyrannosa ...
,'' Summary in press releas
No Olympian: Analysis hints ''Indominus Rex'' ran slowly, if at all
or the flight mechanics of ''Microraptor''. It is relatively commonplace to study the internal details of fossils using X-ray microtomography. Paleontology, biology, archaeology, and
paleoneurobiology Paleoneurobiology is the study of brain evolution by analysis of brain endocasts to determine endocranial traits and volumes. Considered a subdivision of neuroscience, paleoneurobiology combines techniques from other fields of study including pa ...
combine to study endocranial casts (endocasts) of species related to humans to clarify the evolution of the human brain. Paleontology even contributes to
astrobiology Astrobiology, and the related field of exobiology, is an interdisciplinary scientific field that studies the origins, early evolution, distribution, and future of life in the universe. Astrobiology is the multidisciplinary field that invest ...
, the investigation of possible life on other
planet A planet is a large, rounded astronomical body that is neither a star nor its remnant. The best available theory of planet formation is the nebular hypothesis, which posits that an interstellar cloud collapses out of a nebula to create a you ...
s, by developing models of how life may have arisen and by providing techniques for detecting evidence of life.


Subdivisions

As knowledge has increased, paleontology has developed specialised subdivisions.
Vertebrate paleontology Vertebrate paleontology is the subfield of paleontology that seeks to discover, through the study of fossilized remains, the behavior, reproduction and appearance of extinct animals with vertebrae or a notochord. It also tries to connect, by us ...
concentrates on fossils from the earliest fish to the immediate ancestors of modern
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur ...
s.
Invertebrate paleontology Invertebrate paleontology (also spelled invertebrate palaeontology) is sometimes described as invertebrate paleozoology or invertebrate paleobiology. Whether it is considered to be a subfield of paleontology, paleozoology, or paleobiology, this ...
deals with fossils such as
mollusc Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is est ...
s,
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
s,
annelid The annelids (Annelida , from Latin ', "little ring"), also known as the segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecol ...
worms and
echinoderm An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the ...
s.
Paleobotany Paleobotany, which is also spelled as palaeobotany, is the branch of botany dealing with the recovery and identification of plant remains from geological contexts, and their use for the biological reconstruction of past environments (paleogeog ...
studies fossil
plants Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae exclude ...
,
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular micr ...
, and fungi.
Palynology Palynology is the "study of dust" (from grc-gre, παλύνω, palynō, "strew, sprinkle" and ''-logy'') or of "particles that are strewn". A classic palynologist analyses particulate samples collected from the air, from water, or from deposit ...
, the study of
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametop ...
and
spores In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, ...
produced by land plants and
protist A protist () is any eukaryotic organism (that is, an organism whose cells contain a cell nucleus) that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor (the last eukaryotic common ancestor), the e ...
s, straddles paleontology and
botany Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytologist is a scientist who specialises in this field. The term "botany" comes from the Ancient Greek w ...
, as it deals with both living and fossil organisms.
Micropaleontology Micropaleontology (American spelling; spelled micropalaeontology in European usage) is the branch of paleontology (palaeontology) that studies microfossils, or fossils that require the use of a microscope to see the organism, its morphology and it ...
deals with microscopic fossil organisms of all kinds. Instead of focusing on individual organisms, paleoecology examines the interactions between different ancient organisms, such as their
food chain A food chain is a linear network of links in a food web starting from producer organisms (such as grass or algae which produce their own food via photosynthesis) and ending at an apex predator species (like grizzly bears or killer whales), de ...
s, and the two-way interactions with their environments.  For example, the development of
oxygenic photosynthesis Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in ...
by bacteria caused the oxygenation of the atmosphere and hugely increased the productivity and diversity of
ecosystem An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syst ...
s. Together, these led to the evolution of complex
eukaryotic Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
cells, from which all
multicellular A multicellular organism is an organism that consists of more than one cell, in contrast to unicellular organism. All species of animals, land plants and most fungi are multicellular, as are many algae, whereas a few organisms are partially ...
organisms are built.
Paleoclimatology Paleoclimatology (British spelling, palaeoclimatology) is the study of climates for which direct measurements were not taken. As instrumental records only span a tiny part of Earth's history, the reconstruction of ancient climate is important to ...
, although sometimes treated as part of paleoecology, focuses more on the history of Earth's climate and the mechanisms that have changed it – which have sometimes included
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
ary developments, for example the rapid expansion of land plants in the
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, wh ...
period removed more
carbon dioxide Carbon dioxide ( chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is t ...
from the atmosphere, reducing the
greenhouse effect The greenhouse effect is a process that occurs when energy from a planet's host star goes through the planet's atmosphere and heats the planet's surface, but greenhouse gases in the atmosphere prevent some of the heat from returning directly ...
and thus helping to cause an
ice age An ice age is a long period of reduction in the temperature of Earth's surface and atmosphere, resulting in the presence or expansion of continental and polar ice sheets and alpine glaciers. Earth's climate alternates between ice ages and gre ...
in the
Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carboniferou ...
period.
Biostratigraphy Biostratigraphy is the branch of stratigraphy which focuses on correlating and assigning relative ages of rock strata by using the fossil assemblages contained within them.Hine, Robert. “Biostratigraphy.” ''Oxford Reference: Dictionary of ...
, the use of fossils to work out the chronological order in which rocks were formed, is useful to both paleontologists and geologists.
Biogeography Biogeography is the study of the distribution of species and ecosystems in geographic space and through geological time. Organisms and biological communities often vary in a regular fashion along geographic gradients of latitude, elevation, ...
studies the spatial distribution of organisms, and is also linked to geology, which explains how Earth's geography has changed over time.


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. Fossilisation is a rare event, and most fossils are destroyed by
erosion Erosion is the action of surface processes (such as water flow or wind) that removes soil, rock, or dissolved material from one location on the Earth's crust, and then transports it to another location where it is deposited. Erosion is d ...
or
metamorphism Metamorphism is the transformation of existing rock (the protolith) to rock with a different mineral composition or texture. Metamorphism takes place at temperatures in excess of , and often also at elevated pressure or in the presence of ch ...
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. : Non-technica
summary
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. 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 Phyla, the plural of ''phylum'', may refer to: * Phylum, a biological taxon between Kingdom and Class * by analogy, in linguistics, a large division of possibly related languages, or a major language family which is not subordinate to another Phy ...
of living animals, two-thirds have never been found as fossils. Occasionally, unusual environments may preserve soft tissues. These
lagerstätte A Lagerstätte (, from '' Lager'' 'storage, lair' '' Stätte'' 'place'; plural ''Lagerstätten'') is a sedimentary deposit that exhibits extraordinary fossils with exceptional preservation—sometimes including preserved soft tissues. These f ...
n 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. 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.


Trace fossils

Trace fossil A trace fossil, also known as an ichnofossil (; from el, ἴχνος ''ikhnos'' "trace, track"), is a fossil record of biological activity but not the preserved remains of the plant or animal itself. Trace fossils contrast with body fossils, ...
s consist mainly of tracks and burrows, but also include
coprolite A coprolite (also known as a coprolith) is fossilized feces. Coprolites are classified as trace fossils as opposed to body fossils, as they give evidence for the animal's behaviour (in this case, diet) rather than morphology. The name is ...
s (fossil
feces Feces ( or faeces), known colloquially and in slang as poo and poop, are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the large intestine. Feces contain a rela ...
) and marks left by feeding. Trace fossils are particularly significant because they represent a data source that is not limited to animals with easily fossilised 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.e.g. 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
earthworm An earthworm is a terrestrial invertebrate that belongs to the phylum Annelida. They exhibit a tube-within-a-tube body plan; they are externally segmented with corresponding internal segmentation; and they usually have setae on all segments. T ...
s).


Geochemical observations

Geochemical observations may help to deduce the global level of biological activity at a certain period, or the affinity of certain fossils. For example, geochemical features of rocks may reveal when life first arose on Earth, and may provide evidence of the presence of
eukaryotic Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
cells, the type from which all
multicellular A multicellular organism is an organism that consists of more than one cell, in contrast to unicellular organism. All species of animals, land plants and most fungi are multicellular, as are many algae, whereas a few organisms are partially ...
organisms are built. Analyses of
carbon Carbon () is a chemical element with the symbol C and atomic number 6. It is nonmetallic and tetravalent—its atom making four electrons available to form covalent chemical bonds. It belongs to group 14 of the periodic table. Carbon ma ...
isotope ratios may help to explain major transitions such as the
Permian–Triassic extinction event The Permian–Triassic (P–T, P–Tr) extinction event, also known as the Latest Permian extinction event, the End-Permian Extinction and colloquially as the Great Dying, formed the boundary between the Permian and Triassic geologic periods, ...
.


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.
Linnaean taxonomy Linnaean taxonomy can mean either of two related concepts: # The particular form of biological classification (taxonomy) set up by Carl Linnaeus, as set forth in his ''Systema Naturae'' (1735) and subsequent works. In the taxonomy of Linnaeus t ...
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 Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
or
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
or
order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of ...
; this is important since the Linnaean rules for naming groups are tied to their levels, and hence if a group is moved to a different level it must be renamed.
Simple example cladogram
Warm-bloodedness evolved somewhere in the
synapsid–mammal transition.
Warm-bloodedness must also have evolved at one of
these points – an example of
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
.
Paleontologists generally use approaches based on
cladistics Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups (" clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived cha ...
, a technique for working out the evolutionary "family tree" of a set of organisms. 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 Anatomy () is the branch of biology concerned with the study of the structure of organisms and their parts. Anatomy is a branch of natural science that deals with the structural organization of living things. It is an old science, having it ...
, such as the presence of a
notochord In anatomy, the notochord is a flexible rod which is similar in structure to the stiffer cartilage. If a species has a notochord at any stage of its life cycle (along with 4 other features), it is, by definition, a chordate. The notochord consi ...
, or
molecular A molecule is a group of two or more atoms held together by attractive forces known as chemical bonds; depending on context, the term may or may not include ions which satisfy this criterion. In quantum physics, organic chemistry, and bio ...
, by comparing sequences of DNA or
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, res ...
s. 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.
Evolutionary developmental biology Evolutionary developmental biology (informally, evo-devo) is a field of biological research that compares the developmental processes of different organisms to infer how developmental processes evolved. The field grew from 19th-century beginn ...
, commonly abbreviated to "Evo Devo", also helps paleontologists to produce "family trees", and understand fossils. For example, the embryological development of some modern
brachiopod Brachiopods (), phylum Brachiopoda, are a phylum of trochozoan animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, w ...
s suggests that brachiopods may be descendants of the halkieriids, which became extinct in the
Cambrian The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ...
period.


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 Radiometric dating, radioactive dating or radioisotope dating is a technique which is used to date materials such as rocks or carbon, in which trace radioactive impurities were selectively incorporated when they were formed. The method compares ...
. 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. 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. Consequently, paleontologists must usually rely on
stratigraphy Stratigraphy is a branch of geology concerned with the study of rock layers ( strata) and layering (stratification). It is primarily used in the study of sedimentary and layered volcanic rocks. Stratigraphy has three related subfields: lithost ...
to date fossils. Stratigraphy is the science of deciphering the "layer-cake" that is the
sediment Sediment is a naturally occurring material that is broken down by processes of weathering and erosion, and is subsequently transported by the action of wind, water, or ice or by the force of gravity acting on the particles. For example, sand ...
ary record, and has been compared to a
jigsaw puzzle A jigsaw puzzle is a tiling puzzle that requires the assembly of often irregularly shaped interlocking and mosaiced pieces, each of which typically has a portion of a picture. When assembled, the puzzle pieces produce a complete picture. In t ...
. 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. Because rock sequences are not continuous, but may be broken up by faults or periods of
erosion Erosion is the action of surface processes (such as water flow or wind) that removes soil, rock, or dissolved material from one location on the Earth's crust, and then transports it to another location where it is deposited. Erosion is d ...
, 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. If rocks of unknown age are found to have traces of ''E. pseudoplanus'', they must have a mid-Ordovician age. Such
index fossil Biostratigraphy is the branch of stratigraphy which focuses on correlating and assigning relative ages of rock strata by using the fossil assemblages contained within them.Hine, Robert. “Biostratigraphy.” ''Oxford Reference: Dictionary of Bio ...
s 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. 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
continent A continent is any of several large landmasses. Generally identified by convention rather than any strict criteria, up to seven geographical regions are commonly regarded as continents. Ordered from largest in area to smallest, these seven ...
s. 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
mutation In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, m ...
s accumulate at a constant rate. These "
molecular clock The molecular clock is a figurative term for a technique that uses the mutation rate of biomolecules to deduce the time in prehistory when two or more life forms diverged. The biomolecular data used for such calculations are usually nucleo ...
s", 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 The Cambrian explosion, Cambrian radiation, Cambrian diversification, or the Biological Big Bang refers to an interval of time approximately in the Cambrian Period when practically all major animal phyla started appearing in the fossil record. ...
first evolved, and estimates produced by different techniques may vary by a factor of two.


History of life

Earth formed about and, after a collision that formed the
Moon The Moon is Earth's only natural satellite. It is the fifth largest satellite in the Solar System and the largest and most massive relative to its parent planet, with a diameter about one-quarter that of Earth (comparable to the width of ...
about 40 million years later, may have cooled quickly enough to have oceans and an atmosphere about . There is evidence on the Moon of a
Late Heavy Bombardment The Late Heavy Bombardment (LHB), or lunar cataclysm, is a hypothesized event thought to have occurred approximately 4.1 to 3.8 billion years (Ga) ago, at a time corresponding to the Neohadean and Eoarchean eras on Earth. According to the hypot ...
by asteroids from . If, as seems likely, such a bombardment struck Earth at the same time, the first atmosphere and oceans may have been stripped away. Paleontology traces the evolutionary history of life back to over , possibly as far as . The oldest clear evidence of life on Earth dates to , although there have been reports, often disputed, of
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
bacteria from and of geochemical evidence for the presence of life . Some scientists have proposed that life on Earth was "seeded" from elsewhere, but most research concentrates on various explanations of how life could have arisen independently on Earth. For about 2,000 million years microbial mats, multi-layered colonies of different bacteria, were the dominant life on Earth. The evolution of
oxygenic photosynthesis Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in ...
enabled them to play the major role in the oxygenation of the atmosphere from about . This change in the atmosphere increased their effectiveness as nurseries of evolution. While
eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacter ...
s, 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 Poison is a chemical substance that has a detrimental effect to life. The term is used in a wide range of scientific fields and industries, where it is often specifically defined. It may also be applied colloquially or figuratively, with a broa ...
to a powerful source of
metabolic Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run cell ...
energy. This innovation may have come from primitive eukaryotes capturing oxygen-powered bacteria as
endosymbiont An ''endosymbiont'' or ''endobiont'' is any organism that lives within the body or cells of another organism most often, though not always, in a mutualistic relationship. (The term endosymbiosis is from the Greek: ἔνδον ''endon'' "within ...
s and transforming them into
organelle In cell biology, an organelle is a specialized subunit, usually within a cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts of cells, as organs are to the body, hence ''organelle,'' th ...
s called
mitochondria A mitochondrion (; ) is an organelle found in the cells of most Eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is used ...
. The earliest evidence of complex eukaryotes with organelles (such as mitochondria) dates from .
Multicellular A multicellular organism is an organism that consists of more than one cell, in contrast to unicellular organism. All species of animals, land plants and most fungi are multicellular, as are many algae, whereas a few organisms are partially ...
life is composed only of eukaryotic cells, and the earliest evidence for it is the
Francevillian Group Fossil The Francevillian biota (also known as Gabon macrofossils or Gabonionta) is a group of 2.1-billion-year-old Palaeoproterozoic, macroscopic organisms known from fossils found in Gabon in the Palaeoproterozoic Francevillian B Formation, a black s ...
s from , although specialisation of cells for different functions first appears between (a possible fungus) and (a probable
red alga Red algae, or Rhodophyta (, ; ), are one of the oldest groups of eukaryotic algae. The Rhodophyta also comprises one of the largest phyla of algae, containing over 7,000 currently recognized species with taxonomic revisions ongoing. The majority ...
).
Sexual reproduction Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote th ...
may be a prerequisite for specialisation of cells, as an asexual multicellular organism might be at risk of being taken over by rogue cells that retain the ability to reproduce. The earliest known animals are
cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that ...
ns from about , but these are so modern-looking that must be descendants of earlier animals. Early fossils of animals are rare because they had not developed mineralised, easily fossilized hard parts until about . The earliest modern-looking
bilateria The Bilateria or bilaterians are animals with bilateral symmetry as an embryo, i.e. having a left and a right side that are mirror images of each other. This also means they have a head and a tail (anterior-posterior axis) as well as a belly and ...
n animals appear in the Early
Cambrian The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ...
, 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 The Cambrian explosion, Cambrian radiation, Cambrian diversification, or the Biological Big Bang refers to an interval of time approximately in the Cambrian Period when practically all major animal phyla started appearing in the fossil record. ...
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.
Vertebrates Vertebrates () comprise all animal taxa within the subphylum Vertebrata () (chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with ...
remained a minor group until the first jawed fish appeared in the Late
Ordovician The Ordovician ( ) is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start of the Silurian Period Mya. T ...
. The spread of animals and plants from water to land required organisms to solve several problems, including protection against drying out and supporting themselves against
gravity In physics, gravity () is a fundamental interaction which causes mutual attraction between all things with mass or energy. Gravity is, by far, the weakest of the four fundamental interactions, approximately 1038 times weaker than the stro ...
. The earliest evidence of land plants and land invertebrates date back to about and respectively. Those invertebrates, as indicated by their trace and body fossils, were shown to be arthropods known as
euthycarcinoid Euthycarcinoidea are an enigmatic group of extinct possibly amphibious arthropods that ranged from Cambrian to Triassic times. Fossils are known from Europe, North America, Argentina, Australia and Antarctica. Description The euthycarcinoid body ...
s. The lineage that produced land vertebrates evolved later but very rapidly between and ; recent discoveries have overturned earlier ideas about the history and driving forces behind their evolution. Land plants were so successful that their detritus caused an
ecological crisis An ecological or environmental crises occurs when changes to the environment of a species or population destabilizes its continued survival. Some of the important causes include: * Degradation of an abiotic ecological factor (for example, incr ...
in the Late
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, wh ...
, until the evolution of fungi that could digest dead wood. During the
Permian The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleo ...
period,
synapsid Synapsids + (, 'arch') > () "having a fused arch"; synonymous with ''theropsids'' (Greek, "beast-face") are one of the two major groups of animals that evolved from basal amniotes, the other being the sauropsids, the group that includes reptil ...
s, including the ancestors of
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur ...
s, may have dominated land environments, but this ended with the
Permian–Triassic extinction event The Permian–Triassic (P–T, P–Tr) extinction event, also known as the Latest Permian extinction event, the End-Permian Extinction and colloquially as the Great Dying, formed the boundary between the Permian and Triassic geologic periods, ...
, which came very close to wiping out all complex life. The extinctions were apparently fairly sudden, at least among vertebrates. During the slow recovery from this catastrophe a previously obscure group,
archosaur Archosauria () is a clade of diapsids, with birds and crocodilians as the only living representatives. Archosaurs are broadly classified as reptiles, in the cladistic sense of the term which includes birds. Extinct archosaurs include non-avia ...
s, became the most abundant and diverse terrestrial vertebrates. One archosaur group, the dinosaurs, were the dominant land vertebrates for the rest of the
Mesozoic The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretace ...
, and birds evolved from one group of dinosaurs. During this time mammals' ancestors survived only as small, mainly nocturnal
insectivore A robber fly eating a hoverfly An insectivore is a carnivorous animal or plant that eats insects. An alternative term is entomophage, which can also refer to the human practice of eating insects. The first vertebrate insectivores were ...
s, which may have accelerated the development of mammalian traits such as endothermy and hair. After the
Cretaceous–Paleogene extinction event The Cretaceous–Paleogene (K–Pg) extinction event (also known as the Cretaceous–Tertiary extinction) was a sudden mass extinction of three-quarters of the plant and animal species on Earth, approximately 66 million years ago. With the ...
killed off all the dinosaurs except the birds, mammals increased rapidly in size and diversity, and some took to the air and the sea. Fossil evidence indicates that
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants t ...
s appeared and rapidly diversified in the Early
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
between and . Their rapid rise to dominance of terrestrial ecosystems is thought to have been propelled by
coevolution In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well ...
with
pollinating Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds, a ...
insects.
Social insects Eusociality (from Greek εὖ ''eu'' "good" and social), the highest level of organization of sociality, is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping gen ...
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. Humans evolved from a lineage of upright-walking apes whose earliest fossils date from over . 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 . 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 Homininae (), also called "African hominids" or "African apes", is a subfamily of Hominidae. It includes two tribes, with their extant as well as extinct species: 1) the tribe Hominini (with the genus ''Homo'' including modern humans and numerou ...
species, or arose worldwide at the same time as a result of
interbreeding In biology, a hybrid is the offspring resulting from combining the qualities of two organisms of different breeds, varieties, species or genera through sexual reproduction. Hybrids are not always intermediates between their parents (such as in ...
.


Mass extinctions

Life on earth has suffered occasional mass extinctions at least since . Despite their disastrous effects, mass extinctions have sometimes accelerated the evolution of life on earth. When dominance of an
ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...
passes from one group of organisms to another, this is rarely because the new dominant group outcompetes the old, but usually because an extinction event allows a new group, which may posses an advantageous trait, to outlive the old and move into its niche. 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: * The oceans may have become more hospitable to life over the last 500 million years and less vulnerable to mass extinctions:
dissolved oxygen Oxygen saturation (symbol SO2) is a relative measure of the concentration of oxygen that is dissolved or carried in a given medium as a proportion of the maximal concentration that can be dissolved in that medium at the given temperature. It ca ...
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 Eutrophication is the process by which an entire body of water, or parts of it, becomes progressively enriched with minerals and nutrients, particularly nitrogen and phosphorus. It has also been defined as "nutrient-induced increase in phyt ...
and
anoxic event Oceanic anoxic events or anoxic events ( anoxia conditions) describe periods wherein large expanses of Earth's oceans were depleted of dissolved oxygen (O2), creating toxic, euxinic (anoxic and sulfidic) waters. Although anoxic events have not ...
s; marine ecosystems became more diversified so that
food chain A food chain is a linear network of links in a food web starting from producer organisms (such as grass or algae which produce their own food via photosynthesis) and ending at an apex predator species (like grizzly bears or killer whales), de ...
s were less likely to be disrupted. * Reasonably complete
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s 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 Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
, which were often defined solely to accommodate these finds – the story of ''
Anomalocaris ''Anomalocaris'' ("unlike other shrimp", or "abnormal shrimp") is an extinct genus of radiodont, an order of early-diverging stem-group arthropods. The first fossils of ''Anomalocaris'' were discovered in the ''Ogygopsis'' Shale of the Stephen F ...
'' is an example of this. 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 are interpreted as becoming extinct very quickly.
Biodiversity Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic ('' genetic variability''), species ('' species diversity''), and ecosystem ('' ecosystem diversity'') ...
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" shows a different trend: a fairly swift rise from , a slight decline from , in which the devastating
Permian–Triassic extinction event The Permian–Triassic (P–T, P–Tr) extinction event, also known as the Latest Permian extinction event, the End-Permian Extinction and colloquially as the Great Dying, formed the boundary between the Permian and Triassic geologic periods, ...
is an important factor, and a swift rise from to the present.


History

Although paleontology became established around 1800, earlier thinkers had noticed aspects of the
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
record. The ancient Greek
philosopher A philosopher is a person who practices or investigates philosophy. The term ''philosopher'' comes from the grc, φιλόσοφος, , translit=philosophos, meaning 'lover of wisdom'. The coining of the term has been attributed to the Greek th ...
Xenophanes Xenophanes of Colophon (; grc, Ξενοφάνης ὁ Κολοφώνιος ; c. 570 – c. 478 BC) was a Greek philosopher, theologian, poet, and critic of Homer from Ionia who travelled throughout the Greek-speaking world in early Classica ...
(570–480 BCE) concluded from fossil sea shells that some areas of land were once under water. During the
Middle Ages In the history of Europe, the Middle Ages or medieval period lasted approximately from the late 5th to the late 15th centuries, similar to the post-classical period of global history. It began with the fall of the Western Roman Empire ...
the Persian naturalist
Ibn Sina Ibn Sina ( fa, ابن سینا; 980 – June 1037 CE), commonly known in the West as Avicenna (), was a Persian polymath who is regarded as one of the most significant physicians, astronomers, philosophers, and writers of the Islami ...
, known as ''Avicenna'' in Europe, discussed fossils and proposed a theory of petrifying fluids on which
Albert of Saxony en, Frederick Augustus Albert Anthony Ferdinand Joseph Charles Maria Baptist Nepomuk William Xavier George Fidelis , image = Albert of Saxony by Nicola Perscheid c1900.jpg , image_size = , caption = Photograph by Nicola Persch ...
elaborated in the 14th century. The Chinese naturalist
Shen Kuo Shen Kuo (; 1031–1095) or Shen Gua, courtesy name Cunzhong (存中) and pseudonym Mengqi (now usually given as Mengxi) Weng (夢溪翁),Yao (2003), 544. was a Chinese polymathic scientist and statesman of the Song dynasty (960–1279). Shen wa ...
(1031–1095) proposed a theory of climate change based on the presence of
petrified In geology, petrifaction or petrification () is the process by which organic material becomes a fossil through the replacement of the original material and the filling of the original pore spaces with minerals. Petrified wood typifies this p ...
bamboo Bamboos are a diverse group of evergreen perennial flowering plants making up the subfamily Bambusoideae of the grass family Poaceae. Giant bamboos are the largest members of the grass family. The origin of the word "bamboo" is uncertain, ...
in regions that in his time were too dry for bamboo. In
early modern Europe Early modern Europe, also referred to as the post-medieval period, is the period of European history between the end of the Middle Ages and the beginning of the Industrial Revolution, roughly the late 15th century to the late 18th century. Histor ...
, the systematic study of fossils emerged as an integral part of the changes in
natural philosophy Natural philosophy or philosophy of nature (from Latin ''philosophia naturalis'') is the philosophical study of physics, that is, nature and the physical universe. It was dominant before the development of modern science. From the ancien ...
that occurred during the
Age of Reason The Age of reason, or the Enlightenment, was an intellectual and philosophical movement that dominated the world of ideas in Europe during the 17th to 19th centuries. Age of reason or Age of Reason may also refer to: * Age of reason (canon law), ...
. In the Italian Renaissance,
Leonardo da Vinci Leonardo di ser Piero da Vinci (15 April 14522 May 1519) was an Italian polymath of the High Renaissance who was active as a painter, draughtsman, engineer, scientist, theorist, sculptor, and architect. While his fame initially rested on ...
made various significant contributions to the field as well as depicted numerous fossils. Leonardo's contributions are central to the history of paleontology because he established a line of continuity between the two main branches of paleontologyichnology and body fossil paleontology.Baucon, A. (2010). "Leonardo da Vinci, the founding father of ichnology". ''Palaios'' 25. Abstract available from th
author's webpage
/ref> He identified the following: # The biogenic nature of ichnofossils, i.e. ichnofossils were structures left by living organisms; # The utility of ichnofossils as paleoenvironmental toolscertain ichnofossils show the marine origin of rock strata; # The importance of the neoichnological approachrecent traces are a key to understanding ichnofossils; # The independence and complementary evidence of ichnofossils and body fossilsichnofossils are distinct from body fossils, but can be integrated with body fossils to provide paleontological information At the end of the 18th century
Georges Cuvier Jean Léopold Nicolas Frédéric, Baron Cuvier (; 23 August 1769 – 13 May 1832), known as Georges Cuvier, was a French naturalist and zoologist, sometimes referred to as the "founding father of paleontology". Cuvier was a major figure in na ...
's work established
comparative anatomy Comparative anatomy is the study of similarities and differences in the anatomy of different species. It is closely related to evolutionary biology and phylogeny (the evolution of species). The science began in the classical era, continuing in ...
as a scientific discipline and, by proving that some fossil animals resembled no living ones, demonstrated that animals could become
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
, leading to the emergence of paleontology. The expanding knowledge of the fossil record also played an increasing role in the development of geology, particularly
stratigraphy Stratigraphy is a branch of geology concerned with the study of rock layers ( strata) and layering (stratification). It is primarily used in the study of sedimentary and layered volcanic rocks. Stratigraphy has three related subfields: lithost ...
. The first half of the 19th century saw geological and paleontological activity become increasingly well organised with the growth of geologic societies and museums 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. 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 The geologic time scale, or geological time scale, (GTS) is a representation of time based on the rock record of Earth. It is a system of chronological dating that uses chronostratigraphy (the process of relating strata to time) and geochr ...
, largely based on fossil evidence. Although she was rarely recognised by the scientific community, Mary Anning was a significant contributor to the field of palaeontology during this period; she uncovered multiple novel
Mesozoic The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretace ...
reptile fossils and deducted that what were then known as
bezoar A bezoar is a mass often found trapped in the gastrointestinal system, though it can occur in other locations. A pseudobezoar is an indigestible object introduced intentionally into the digestive system. There are several varieties of bezoar, s ...
stones are in fact fossilised faeces. In 1822 Henri Marie Ducrotay de Blainville, editor of ''Journal de Physique'', coined the word "palaeontology" to refer to the study of ancient living organisms through fossils. 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 Transmutation of species and transformism are unproven 18th and 19th-century evolutionary ideas about the change of one species into another that preceded Charles Darwin's theory of natural selection. The French ''Transformisme'' was a term used ...
. After
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended ...
published '' Origin of Species'' in 1859, much of the focus of paleontology shifted to understanding
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
ary paths, including
human evolution Human evolution is the evolutionary process within the history of primates that led to the emergence of '' Homo sapiens'' as a distinct species of the hominid family, which includes the great apes. This process involved the gradual developmen ...
, and evolutionary theory. 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. 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. The last few decades of the 20th century saw a renewed interest in
mass extinction An extinction event (also known as a mass extinction or biotic crisis) is a widespread and rapid decrease in the biodiversity on Earth. Such an event is identified by a sharp change in the diversity and abundance of multicellular organisms. I ...
s and their role in the evolution of life on Earth. There was also a renewed interest in the
Cambrian explosion The Cambrian explosion, Cambrian radiation, Cambrian diversification, or the Biological Big Bang refers to an interval of time approximately in the Cambrian Period when practically all major animal phyla started appearing in the fossil record. ...
that apparently saw the development of the body plans of most animal
phyla Phyla, the plural of ''phylum'', may refer to: * Phylum, a biological taxon between Kingdom and Class * by analogy, in linguistics, a large division of possibly related languages, or a major language family which is not subordinate to another Phy ...
. The discovery of fossils of the
Ediacaran biota The Ediacaran (; formerly Vendian) biota is a taxonomic period classification that consists of all life forms that were present on Earth during the Ediacaran Period (). These were composed of enigmatic tubular and frond-shaped, mostly sess ...
and developments in
paleobiology Paleobiology (or palaeobiology) is an interdisciplinary field that combines the methods and findings found in both the earth sciences and the life sciences. Paleobiology is not to be confused with geobiology, which focuses more on the interactio ...
extended knowledge about the history of life back far before the Cambrian. Increasing awareness of
Gregor Mendel Gregor Johann Mendel, OSA (; cs, Řehoř Jan Mendel; 20 July 1822 – 6 January 1884) was a biologist, meteorologist, mathematician, Augustinian friar and abbot of St. Thomas' Abbey in Brünn (''Brno''), Margraviate of Moravia. Mendel was ...
's pioneering work in
genetics Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinian friar work ...
led first to the development of
population genetics Population genetics is a subfield of genetics that deals with genetic differences within and between populations, and is a part of evolutionary biology. Studies in this branch of biology examine such phenomena as adaptation, speciation, and po ...
and then in the mid-20th century to the modern evolutionary synthesis, which explains
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
as the outcome of events such as
mutation In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, m ...
s and
horizontal gene transfer Horizontal gene transfer (HGT) or lateral gene transfer (LGT) is the movement of genetic material between unicellular and/or multicellular organisms other than by the ("vertical") transmission of DNA from parent to offspring (reproduction). H ...
, which provide
genetic variation Genetic variation is the difference in DNA among individuals or the differences between populations. The multiple sources of genetic variation include mutation and genetic recombination. Mutations are the ultimate sources of genetic variation, b ...
, with
genetic drift Genetic drift, also known as allelic drift or the Wright effect, is the change in the frequency of an existing gene variant (allele) in a population due to random chance. Genetic drift may cause gene variants to disappear completely and there ...
and
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Cha ...
driving changes in this variation over time. 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. In the 1960s
molecular phylogenetics Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
, the investigation of evolutionary "family trees" by techniques derived from
biochemistry Biochemistry or biological chemistry is the study of chemical processes within and relating to living organisms. A sub-discipline of both chemistry and biology, biochemistry may be divided into three fields: structural biology, enzymology and ...
, 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. Although this early study compared
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, res ...
s from apes and humans, most molecular phylogenetics research is now based on comparisons of RNA and DNA.


Paleontology in the vernacular press

Books catered to the general public on paleontology include: * The Last Days of the Dinosaurs: An Asteroid Extinction, and the Beginning of our World written by Riley Black * The Rise and Reign of the Mammals: A New History, from the Shadow of the Dinosaurs to Uswritten by Steve Brusatte * Otherlands: A Journey Through Earth's Extinct Worlds written by Thomas Halliday


See also

* * * * (''with link directory'') * List of notable fossils *
List of paleontologists This is a list of notable paleontologists who have made significant contributions to the field of paleontology. Only paleontologists with biographical articles in Wikipedia are listed here. A * Othenio Abel (Australia, 1875-1946) * William ...
* List of transitional fossils * * * * * * * * * ''
Une Femme ou Deux ''Une Femme ou Deux'' (English: ''One Woman or Two'') is a French screwball comedy romance film released in 1985. It was directed by , who was also the screenwriter along with Élisabeth Rappeneau. It stars Gérard Depardieu, Sigourney Weaver, ...
'' -
French French (french: français(e), link=no) may refer to: * Something of, from, or related to France ** French language, which originated in France, and its various dialects and accents ** French people, a nation and ethnic group identified with Franc ...
screwball comedy Screwball comedy is a subgenre of the romantic comedy genre that became popular during the Great Depression, beginning in the early 1930s and thriving until the early 1940s, that satirizes the traditional love story. It has secondary characteristi ...
romance film starring
Gérard Depardieu Gérard Xavier Marcel Depardieu, CQ (, , ; born 27 December 1948) is a French actor, filmmaker, businessman and vineyard owner since 1989 who is one of the most prolific thespians in film history having completed over 250 films since 1967 al ...
as a paleontologist.


Notes


References


External links


Smithsonian's Paleobiology website

University of California Museum of Paleontology

The Paleontological Society

The Palaeontological Association

The Society of Vertebrate Paleontology

The Paleontology Portal

"Geology, Paleontology & Theories of the Earth"
– a collection of more than 100 digitised landmark and early books on Earth sciences at the
Linda Hall Library The Linda Hall Library is a privately endowed American library of science, engineering and technology located in Kansas City, Missouri, sitting "majestically on a urban arboretum." It is the "largest independently funded public library of scien ...
{{Good article Earth sciences Evolutionary biology * Historical geology