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A fossil (from
Classical Latin Classical Latin is the form of Literary Latin recognized as a literary standard by writers of the late Roman Republic and early Roman Empire. It was used from 75 BC to the 3rd century AD, when it developed into Late Latin. In some later period ...
, ) is any preserved remains, impression, or trace of any once-living thing from a past
geological age 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 geochronol ...
. Examples include
bone A bone is a rigid organ that constitutes part of the skeleton in most vertebrate animals. Bones protect the various other organs of the body, produce red and white blood cells, store minerals, provide structure and support for the body, ...
s, shells,
exoskeletons An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton (endoskeleton) in for example, a human. In usage, some of the ...
, stone imprints of animals or
microbes A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in olde ...
, objects preserved in
amber Amber is fossilized tree resin that has been appreciated for its color and natural beauty since Neolithic times. Much valued from antiquity to the present as a gemstone, amber is made into a variety of decorative objects."Amber" (2004). In ...
, hair,
petrified wood Petrified wood, also known as petrified tree (from Ancient Greek meaning 'rock' or 'stone'; literally 'wood turned into stone'), is the name given to a special type of '' fossilized wood'', the fossilized remains of terrestrial vegetation. ' ...
and DNA remnants. The totality of fossils is known as the ''fossil record''.
Paleontology Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
is the study of fossils: their age, method of formation, and
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 significance. Specimens are usually considered to be fossils if they are over 10,000 years old. The oldest fossils are around 3.48 billion years old to 4.1 billion years old. Early edition, published online before print. The observation in the 19th century that certain fossils were associated with certain rock strata led to the recognition of a
geological timescale 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 geochron ...
and the relative ages of different fossils. The development of
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 ...
techniques in the early 20th century allowed scientists to quantitatively measure the absolute ages of rocks and the fossils they host. There are many processes that lead to fossilization, including
permineralization Permineralization is a process of fossilization of bones and tissues in which mineral deposits form internal casts of organisms. Carried by water, these minerals fill the spaces within organic tissue. Because of the nature of the casts, perminera ...
, casts and molds, authigenic mineralization, replacement and recrystallization, adpression,
carbonization Carbonization is the conversion of organic matters like plants and dead animal remains into carbon through destructive distillation. Complexity in carbonization Carbonization is a pyrolytic reaction, therefore, is considered a complex proces ...
, and bioimmuration. Fossils vary in size from one-
micrometre The micrometre ( international spelling as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer (American spelling), also commonly known as a micron, is a unit of length in the International System of Unit ...
(1 µm) bacteria to
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
s and trees, many meters long and weighing many tons. A fossil normally preserves only a portion of the deceased organism, usually that portion that was partially mineralized during life, such as the bones and teeth of
vertebrate 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 c ...
s, or the chitinous or
calcareous Calcareous () is an adjective meaning "mostly or partly composed of calcium carbonate", in other words, containing lime or being chalky. The term is used in a wide variety of scientific disciplines. In zoology ''Calcareous'' is used as an ad ...
exoskeletons of
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
s. Fossils may also consist of the marks left behind by the organism while it was alive, such as animal tracks or feces (
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 de ...
s). These types of fossil are called
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 or ''ichnofossils'', as opposed to ''body fossils''. Some fossils are biochemical and are called ''chemofossils'' or biosignatures.


Reliability

Though the fossil record is incomplete, numerous studies have demonstrated that there is enough information available to give us a good understanding of the pattern of diversification of life on Earth. In addition, the record can predict and fill gaps such as the discovery of
Tiktaalik ''Tiktaalik'' (; Inuktitut ) is a monospecific genus of extinct sarcopterygian (lobe-finned fish) from the Late Devonian Period, about 375 Mya (million years ago), having many features akin to those of tetrapods (four-legged animals). It may ha ...
in the arctic of
Canada Canada is a country in North America. Its ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, covering over , making it the world's second-largest country by tot ...
.


Fossilization processes

The process of fossilization varies according to tissue type and external conditions.


Permineralization

Permineralization Permineralization is a process of fossilization of bones and tissues in which mineral deposits form internal casts of organisms. Carried by water, these minerals fill the spaces within organic tissue. Because of the nature of the casts, perminera ...
is a process of fossilization that occurs when an organism is buried. The empty spaces within an organism (spaces filled with liquid or gas during life) become filled with mineral-rich
groundwater Groundwater is the water present beneath Earth's surface in rock and soil pore spaces and in the fractures of rock formations. About 30 percent of all readily available freshwater in the world is groundwater. A unit of rock or an unconsolidated ...
. Minerals precipitate from the groundwater, occupying the empty spaces. This process can occur in very small spaces, such as within the cell wall of a plant cell. Small scale permineralization can produce very detailed fossils. For permineralization to occur, the organism must become covered by sediment soon after death, otherwise the remains are destroyed by scavengers or decomposition. The degree to which the remains are decayed when covered determines the later details of the fossil. Some fossils consist only of skeletal remains or teeth; other fossils contain traces of
skin Skin is the layer of usually soft, flexible outer tissue covering the body of a vertebrate animal, with three main functions: protection, regulation, and sensation. Other animal coverings, such as the arthropod exoskeleton, have different de ...
, feathers or even soft tissues. This is a form of
diagenesis Diagenesis () is the process that describes physical and chemical changes in sediments first caused by water-rock interactions, microbial activity, and compaction after their deposition. Increased pressure and temperature only start to play a ...
.


Casts and molds

In some cases, the original remains of the organism completely dissolve or are otherwise destroyed. The remaining organism-shaped hole in the rock is called an ''external mold''. If this void is later filled with sediment, the resulting ''cast'' resembles what the organism looked like. An
endocast An endocast is the internal cast of a hollow object, often referring to the cranial vault in the study of brain development in humans and other organisms. Endocasts can be artificially made for examining the properties of a hollow, inaccessible sp ...
, or ''internal mold'', is the result of sediments filling an organism's interior, such as the inside of a bivalve or
snail A snail is, in loose terms, a shelled gastropod. The name is most often applied to land snails, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class G ...
or the hollow of a
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
. Endocasts are sometimes termed , especially when bivalves are preserved this way.


Authigenic mineralization

This is a special form of cast and mold formation. If the chemistry is right, the organism (or fragment of organism) can act as a nucleus for the precipitation of minerals such as
siderite Siderite is a mineral composed of iron(II) carbonate (FeCO3). It takes its name from the Greek word σίδηρος ''sideros,'' "iron". It is a valuable iron mineral, since it is 48% iron and contains no sulfur or phosphorus. Zinc, magnesium and ...
, resulting in a nodule forming around it. If this happens rapidly before significant decay to the organic tissue, very fine three-dimensional morphological detail can be preserved. Nodules from the Carboniferous
Mazon Creek fossil beds The Mazon Creek fossil beds are a conservation ' found near Morris, in Grundy County, Illinois. The fossils are preserved in ironstone concretions, formed approximately in the mid- Pennsylvanian epoch of the Carboniferous period. These concret ...
of Illinois, USA, are among the best documented examples of such mineralization.


Replacement and recrystallization

Replacement occurs when the shell, bone, or other tissue is replaced with another mineral. In some cases mineral replacement of the original shell occurs so gradually and at such fine scales that microstructural features are preserved despite the total loss of original material. A shell is said to be ''recrystallized'' when the original skeletal compounds are still present but in a different crystal form, as from
aragonite Aragonite is a carbonate mineral, one of the three most common naturally occurring crystal forms of calcium carbonate, (the other forms being the minerals calcite and vaterite). It is formed by biological and physical processes, including pre ...
to calcite.


Adpression (compression-impression)

Compression fossil A compression fossil is a fossil preserved in sedimentary rock that has undergone physical compression. While it is uncommon to find animals preserved as good compression fossils, it is very common to find plants preserved this way. The reason f ...
s, such as those of fossil ferns, are the result of chemical reduction of the complex organic molecules composing the organism's tissues. In this case the fossil consists of original material, albeit in a geochemically altered state. This chemical change is an expression of
diagenesis Diagenesis () is the process that describes physical and chemical changes in sediments first caused by water-rock interactions, microbial activity, and compaction after their deposition. Increased pressure and temperature only start to play a ...
. Often what remains is a
carbonaceous film A carbonaceous film or carbon film is an organism outline of a fossil. It is a type of fossil found in any rock when organic material is compressed, leaving only a carbon residue or film. When an organism is buried under many layers of sedimen ...
known as a phytoleim, in which case the fossil is known as a compression. Often, however, the phytoleim is lost and all that remains is an impression of the organism in the rock—an impression fossil. In many cases, however, compressions and impressions occur together. For instance, when the rock is broken open, the phytoleim will often be attached to one part (compression), whereas the counterpart will just be an impression. For this reason, one term covers the two modes of preservation: ''adpression''.


Soft tissue, cell and molecular preservation

Because of their antiquity, an unexpected exception to the alteration of an organism's tissues by chemical reduction of the complex organic molecules during fossilization has been the discovery of soft tissue in dinosaur fossils, including blood vessels, and the isolation of proteins and evidence for DNA fragments. In 2014, Mary Schweitzer and her colleagues reported the presence of iron particles (
goethite Goethite (, ) is a mineral of the diaspore group, consisting of iron(III) oxide-hydroxide, specifically the "α" polymorph. It is found in soil and other low-temperature environments such as sediment. Goethite has been well known since ancient t ...
-aFeO(OH)) associated with soft tissues recovered from dinosaur fossils. Based on various experiments that studied the interaction of iron in
haemoglobin Hemoglobin (haemoglobin BrE) (from the Greek word αἷμα, ''haîma'' 'blood' + Latin ''globus'' 'ball, sphere' + ''-in'') (), abbreviated Hb or Hgb, is the iron-containing oxygen-transport metalloprotein present in red blood cells (erythrocyte ...
with blood vessel tissue they proposed that solution hypoxia coupled with iron
chelation Chelation is a type of bonding of ions and molecules to metal ions. It involves the formation or presence of two or more separate coordinate bonds between a polydentate (multiple bonded) ligand and a single central metal atom. These ligands are ...
enhances the stability and preservation of soft tissue and provides the basis for an explanation for the unforeseen preservation of fossil soft tissues. However, a slightly older study based on eight taxa ranging in time from the Devonian to the
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
found that reasonably well-preserved fibrils that probably represent collagen were preserved in all these fossils and that the quality of preservation depended mostly on the arrangement of the collagen fibers, with tight packing favoring good preservation. There seemed to be no correlation between geological age and quality of preservation, within that timeframe.


Carbonization and coalification

Fossils that are carbonized or coalified consist of the organic remains which have been reduced primarily to the chemical element carbon. Carbonized fossils consist of a thin film which forms a silhouette of the original organism, and the original organic remains were typically soft tissues. Coalified fossils consist primarily of coal, and the original organic remains were typically woody in composition. File:Probable leech from the Waukesha Biota.jpg, Carbonized fossil of a possible leech from the Silurian
Waukesha Biota The Waukesha Biota (also known as Waukesha Lagerstätte, Brandon Bridge Lagerstätte, or Brandon Bridge fauna) refers to a Konservat-Lagerstätte ( a fossil site that preserves soft bodied remains) of Early Silurian ( Telychian to Sheinwoodian) ...
of Wisconsin. File:Lycopod axis.jpg, Partially coalified axis (branch) of a
lycopod Lycopodiopsida is a class of vascular plants known as lycopods, lycophytes or other terms including the component lyco-. Members of the class are also called clubmosses, firmosses, spikemosses and quillworts. They have dichotomously branching s ...
from the Devonian of
Wisconsin Wisconsin () is a state in the upper Midwestern United States. Wisconsin is the 25th-largest state by total area and the 20th-most populous. It is bordered by Minnesota to the west, Iowa to the southwest, Illinois to the south, Lake M ...
.


Bioimmuration

Bioimmuration occurs when a skeletal organism overgrows or otherwise subsumes another organism, preserving the latter, or an impression of it, within the skeleton. Usually it is a
sessile Sessility, or sessile, may refer to: * Sessility (motility), organisms which are not able to move about * Sessility (botany), flowers or leaves that grow directly from the stem or peduncle of a plant * Sessility (medicine), tumors and polyps that ...
skeletal organism, such as a bryozoan or an
oyster Oyster is the common name for a number of different families of salt-water bivalve molluscs that live in marine or brackish habitats. In some species, the valves are highly calcified, and many are somewhat irregular in shape. Many, but not ...
, which grows along a substrate, covering other sessile sclerobionts. Sometimes the bioimmured organism is soft-bodied and is then preserved in negative relief as a kind of external mold. There are also cases where an organism settles on top of a living skeletal organism that grows upwards, preserving the settler in its skeleton. Bioimmuration is known in the fossil record from the
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 ...
to the Recent.


Types


Index

Index fossils (also known as guide fossils, indicator fossils or zone fossils) are fossils used to define and identify geologic periods (or faunal stages). They work on the premise that, although different
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, sa ...
s may look different depending on the conditions under which they were deposited, they may include the remains of the same
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of fossil. The shorter the species' time range, the more precisely different sediments can be correlated, and so rapidly evolving species' fossils are particularly valuable. The best index fossils are common, easy to identify at species level and have a broad distribution—otherwise the likelihood of finding and recognizing one in the two sediments is poor.


Trace

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 de ...
s (fossil feces) and marks left by feeding. 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 animal behaviours. 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). 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. They were first described by
William Buckland William Buckland DD, FRS (12 March 1784 – 14 August 1856) was an English theologian who became Dean of Westminster. He was also a geologist and palaeontologist. Buckland wrote the first full account of a fossil dinosaur, which he named ' ...
in 1829. Prior to this they were known as "fossil fir cones" and "
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." They serve a valuable purpose in paleontology because they provide direct evidence of the predation and diet of extinct organisms. Coprolites may range in size from a few millimetres to over 60 centimetres. File:CambrianRusophycus.jpg, Cambrian
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 including '' Rusophycus'', made by a
trilobite Trilobites (; meaning "three lobes") are extinct marine arthropods that form the class Trilobita. Trilobites form one of the earliest-known groups of arthropods. The first appearance of trilobites in the fossil record defines the base of the ...
File:Coprolite.jpg, A coprolite of a carnivorous dinosaur found in southwestern
Saskatchewan Saskatchewan ( ; ) is a province in western Canada, bordered on the west by Alberta, on the north by the Northwest Territories, on the east by Manitoba, to the northeast by Nunavut, and on the south by the U.S. states of Montana and North Dak ...
File:Climactichnites wilsoni, densely packed.jpg, Densely packed, subaerial or nearshore trackways (''
Climactichnites ''Climactichnites'' is an enigmatic, Cambrian fossil formed on or within sandy tidal flats around . It has been interpreted in many different ways in the past, but is now thought to be a trace fossil of a slug-like organism that moved by crawling ...
wilsoni'') made by a putative, slug-like
mollusk 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 e ...
on a Cambrian tidal flat


Transitional

A ''transitional fossil'' is any fossilized remains of a life form that exhibits traits common to both an ancestral group and its derived descendant group. This is especially important where the descendant group is sharply differentiated by gross anatomy and mode of living from the ancestral group. Because of the incompleteness of the fossil record, there is usually no way to know exactly how close a transitional fossil is to the point of divergence. These fossils serve as a reminder that taxonomic divisions are human constructs that have been imposed in hindsight on a continuum of variation.


Microfossils

Microfossil A microfossil is a fossil that is generally between 0.001 mm and 1 mm in size, the visual study of which requires the use of light or electron microscopy. A fossil which can be studied with the naked eye or low-powered magnification, ...
is a descriptive term applied to fossilized plants and animals whose size is just at or below the level at which the fossil can be analyzed by the naked eye. A commonly applied cutoff point between "micro" and "macro" fossils is 1 mm. Microfossils may either be complete (or near-complete) organisms in themselves (such as the marine plankters
foraminifera Foraminifera (; Latin for "hole bearers"; informally called "forams") are single-celled organisms, members of a phylum or class of amoeboid protists characterized by streaming granular ectoplasm for catching food and other uses; and commonly ...
and
coccolithophore Coccolithophores, or coccolithophorids, are single celled organisms which are part of the phytoplankton, the autotrophic (self-feeding) component of the plankton community. They form a group of about 200 species, and belong either to the king ...
s) or component parts (such as small teeth or
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, ...
) of larger animals or plants. Microfossils are of critical importance as a reservoir of paleoclimate information, and are also commonly used by biostratigraphers to assist in the correlation of rock units.


Resin

Fossil resin (colloquially called
amber Amber is fossilized tree resin that has been appreciated for its color and natural beauty since Neolithic times. Much valued from antiquity to the present as a gemstone, amber is made into a variety of decorative objects."Amber" (2004). In ...
) is a natural
polymer A polymer (; Greek '' poly-'', "many" + ''-mer'', "part") is a substance or material consisting of very large molecules called macromolecules, composed of many repeating subunits. Due to their broad spectrum of properties, both synthetic a ...
found in many types of strata throughout the world, even the
Arctic The Arctic ( or ) is a polar region located at the northernmost part of Earth. The Arctic consists of the Arctic Ocean, adjacent seas, and parts of Canada (Yukon, Northwest Territories, Nunavut), Danish Realm (Greenland), Finland, Iceland, N ...
. The oldest fossil resin dates to the
Triassic The Triassic ( ) is a geologic period and system (stratigraphy), system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago (Year#Abbreviations yr and ya, Mya), to the beginning of the Jurassic Period 201.36 ...
, though most dates to the Cenozoic. The excretion of the resin by certain plants is thought to be an evolutionary adaptation for protection from insects and to seal wounds. Fossil resin often contains other fossils called inclusions that were captured by the sticky resin. These include bacteria, fungi, other plants, and animals. Animal inclusions are usually small
invertebrates Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordat ...
, predominantly
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 such as insects and spiders, and only extremely rarely a
vertebrate 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 c ...
such as a small lizard. Preservation of inclusions can be exquisite, including small fragments of DNA.


Derived, or reworked

A ''derived'', ''reworked'' or is a fossil found in rock that accumulated significantly later than when the fossilized animal or plant died. Reworked fossils are created by erosion exhuming (freeing) fossils from the rock formation in which they were originally deposited and their redeposition in a younger sedimentary deposit.


Wood

Fossil wood is wood that is preserved in the fossil record. Wood is usually the part of a plant that is best preserved (and most easily found). Fossil wood may or may not be
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 proce ...
. The fossil wood may be the only part of the plant that has been preserved: therefore such wood may get a special kind of
botanical name A botanical name is a formal scientific name conforming to the ''International Code of Nomenclature for algae, fungi, and plants'' (ICN) and, if it concerns a plant cultigen, the additional cultivar or Group epithets must conform to the ''Internat ...
. This will usually include "xylon" and a term indicating its presumed affinity, such as ''
Araucarioxylon ''Araucarioxylon arizonicum'' is an extinct species of conifer that is the state fossil of Arizona. The species is known from massive tree trunks that weather out of the Chinle Formation in desert badlands of northern Arizona and adjacent New Mex ...
'' (wood of '' Araucaria'' or some related genus), '' Palmoxylon'' (wood of an indeterminate
palm Palm most commonly refers to: * Palm of the hand, the central region of the front of the hand * Palm plants, of family Arecaceae **List of Arecaceae genera * Several other plants known as "palm" Palm or Palms may also refer to: Music * Palm (ba ...
), or ''Castanoxylon'' (wood of an indeterminate chinkapin).


Subfossil

The term subfossil can be used to refer to remains, such as bones, nests, or fecal deposits, whose fossilization process is not complete, either because the length of time since the animal involved was living is too short (less than 10,000 years) or because the conditions in which the remains were buried were not optimal for fossilization. Subfossils are often found in caves or other shelters where they can be preserved for thousands of years. The main importance of subfossil vs. fossil remains is that the former contain organic material, which can be used for radiocarbon dating or extraction and sequencing of DNA,
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 ...
, or other biomolecules. Additionally,
isotope Isotopes are two or more types of atoms that have the same atomic number (number of protons in their nuclei) and position in the periodic table (and hence belong to the same chemical element), and that differ in nucleon numbers (mass numb ...
ratios can provide much information about the ecological conditions under which extinct animals lived. Subfossils are useful for studying the evolutionary history of an environment and can be important to studies in
paleoclimatology Paleoclimatology (American and British English spelling differences, 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 ...
. Subfossils are often found in depositionary environments, such as lake sediments, oceanic sediments, and soils. Once deposited, physical and chemical
weathering Weathering is the deterioration of rocks, soils and minerals as well as wood and artificial materials through contact with water, atmospheric gases, and biological organisms. Weathering occurs ''in situ'' (on site, with little or no movement) ...
can alter the state of preservation, and small subfossils can also be ingested by living
organisms In biology, an organism () is any life, living system that functions as an individual entity. All organisms are composed of cells (cell theory). Organisms are classified by taxonomy (biology), taxonomy into groups such as Multicellular o ...
. Subfossil remains that date from 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 ...
are exceptionally rare, are usually in an advanced state of decay, and are consequently much disputed. The vast bulk of subfossil material comes from Quaternary sediments, including many subfossilized
chironomid The Chironomidae (informally known as chironomids, nonbiting midges, or lake flies) comprise a family of nematoceran flies with a global distribution. They are closely related to the Ceratopogonidae, Simuliidae, and Thaumaleidae. Many specie ...
head capsules,
ostracod Ostracods, or ostracodes, are a class of the Crustacea (class Ostracoda), sometimes known as seed shrimp. Some 70,000 species (only 13,000 of which are extant) have been identified, grouped into several orders. They are small crustaceans, typi ...
carapaces,
diatoms A diatom (New Latin, Neo-Latin ''diatoma''), "a cutting through, a severance", from el, διάτομος, diátomos, "cut in half, divided equally" from el, διατέμνω, diatémno, "to cut in twain". is any member of a large group com ...
, and
foraminifera Foraminifera (; Latin for "hole bearers"; informally called "forams") are single-celled organisms, members of a phylum or class of amoeboid protists characterized by streaming granular ectoplasm for catching food and other uses; and commonly ...
. For remains such as molluscan seashells, which frequently do not change their chemical composition over geological time, and may occasionally even retain such features as the original color markings for millions of years, the label 'subfossil' is applied to shells that are understood to be thousands of years old, but are of
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 ...
age, and therefore are not old enough to be from the
Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was finally confirmed in ...
epoch.


Chemical fossils

Chemical fossils, or chemofossils, are chemicals found in rocks and fossil fuels (petroleum, coal, and natural gas) that provide an organic signature for ancient life. Molecular fossils and isotope ratios represent two types of chemical fossils. The oldest traces of life on Earth are fossils of this type, including carbon isotope anomalies found in
zircon Zircon () is a mineral belonging to the group of nesosilicates and is a source of the metal zirconium. Its chemical name is zirconium(IV) silicate, and its corresponding chemical formula is Zr SiO4. An empirical formula showing some of t ...
s that imply the existence of life as early as 4.1 billion years ago.


Dating


Estimating dates

Paleontology seeks to map out how life evolved across geologic time. A substantial hurdle is the difficulty of working out fossil ages. 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 careful laboratory work, its basic principle is simple: the rates at which various radioactive elements
decay Decay may refer to: Science and technology * Bit decay, in computing * Software decay, in computing * Distance decay, in geography * Decay time (fall time), in electronics Biology * Decomposition of organic matter * Tooth decay (dental caries ...
are known, and so the ratio of the radioactive element to its decay products 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 volcanic ash layers, which may provide termini for the intervening sediments.


Stratigraphy

Consequently, palaeontologists rely on stratigraphy 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, sa ...
ary record. 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 is claimed to 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 dis ...
, it is very difficult to match up rock beds that are not directly adjacent. However, fossils of species that survived for a relatively short time can be used to match 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 have traces of ''E. pseudoplanus'', they 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 occupy a short time range to be useful. Misleading results are produced if the index fossils are incorrectly dated. 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 rocks of the same age across
continent A continent is any of several large landmasses. Generally identified by convention rather than any strict criteria, up to seven geographical regions In geography, regions, otherwise referred to as zones, lands or territories, are areas t ...
s. Family-tree relationships 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 earlier. It is also possible to estimate how long ago two living clades diverged, in other words 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 replication, DNA or viral repl ...
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 approximate timing: for example, they are not sufficiently precise and reliable for estimating when the groups that feature in the Cambrian explosion first evolved, and estimates produced by different techniques may vary by a factor of two.


Limitations

Organisms are only rarely preserved as fossils in the best of circumstances, and only a fraction of such fossils have been discovered. This is illustrated by the fact that the number of species known through the fossil record is less than 5% of the number of known living species, suggesting that the number of species known through fossils must be far less than 1% of all the species that have ever lived. Because of the specialized and rare circumstances required for a biological structure to fossilize, only a small percentage of life-forms can be expected to be represented in discoveries, and each discovery represents only a snapshot of the process of evolution. The transition itself can only be illustrated and corroborated by transitional fossils, which will never demonstrate an exact half-way point. The fossil record is strongly biased toward organisms with hard-parts, leaving most groups of soft-bodied organisms with little to no role. It is replete with the mollusks, the
vertebrate 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 c ...
s, the
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 ...
s, the
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 and some groups of
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.


Sites


Lagerstätten

Fossil sites with exceptional preservation—sometimes including preserved soft tissues—are known as
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—German for "storage places". These formations may have resulted from carcass burial in an
anoxic The term anoxia means a total depletion in the level of oxygen, an extreme form of hypoxia or "low oxygen". The terms anoxia and hypoxia are used in various contexts: * Anoxic waters, sea water, fresh water or groundwater that are depleted of diss ...
environment with minimal bacteria, thus slowing decomposition. Lagerstätten span geological time from the Cambrian period to the present. Worldwide, some of the best examples of near-perfect fossilization are the Cambrian
Maotianshan shales The Maotianshan Shales are a series of Early Cambrian deposits in the Chiungchussu Formation, famous for their '' Konservat Lagerstätten'', deposits known for the exceptional preservation of fossilized organisms or traces. The Maotianshan Shales ...
and
Burgess Shale The Burgess Shale is a fossil-bearing deposit exposed in the Canadian Rockies of British Columbia, Canada. It is famous for the exceptional preservation of the soft parts of its fossils. At old (middle Cambrian), it is one of the earliest fo ...
, the Devonian Hunsrück Slates, the
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
Solnhofen limestone, and the Carboniferous
Mazon Creek The Mazon Creek fossil beds are a conservation ' found near Morris, in Grundy County, Illinois. The fossils are preserved in ironstone concretions, formed approximately in the mid- Pennsylvanian epoch of the Carboniferous period. These concreti ...
localities.


Stromatolites

Stromatolites are layered accretionary structures formed in shallow water by the trapping, binding and cementation of sedimentary grains by
biofilm A biofilm comprises any syntrophic consortium of microorganisms in which cells stick to each other and often also to a surface. These adherent cells become embedded within a slimy extracellular matrix that is composed of extracellular ...
s of
microorganism A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in olde ...
s, especially cyanobacteria. Stromatolites provide some of the most ancient fossil records of life on Earth, dating back more than 3.5 billion years ago. Stromatolites were much more abundant in Precambrian times. While older,
Archean The Archean Eon ( , also spelled Archaean or Archæan) is the second of four geologic eons of Earth's history, representing the time from . The Archean was preceded by the Hadean Eon and followed by the Proterozoic. The Earth during the Arc ...
fossil remains are presumed to be
colonies In modern parlance, a colony is a territory subject to a form of foreign rule. Though dominated by the foreign colonizers, colonies remain separate from the administration of the original country of the colonizers, the '' metropolitan state'' ...
of cyanobacteria, younger (that is, Proterozoic) fossils may be primordial forms of the eukaryote chlorophytes (that is, green algae). One
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 ...
of stromatolite very common in the
geologic record The geologic record in stratigraphy, paleontology and other natural sciences refers to the entirety of the layers of rock strata. That is, deposits laid down by volcanism or by deposition of sediment derived from weathering detritus (clays, sand ...
is '' Collenia''. The earliest stromatolite of confirmed microbial origin dates to 2.724 billion years ago. A 2009 discovery provides strong evidence of microbial stromatolites extending as far back as 3.45 billion years ago. Stromatolites are a major constituent of the fossil record for life's first 3.5 billion years, peaking about 1.25 billion years ago. They subsequently declined in abundance and diversity, which by the start of the Cambrian had fallen to 20% of their peak. The most widely supported explanation is that stromatolite builders fell victims to grazing creatures (the
Cambrian substrate revolution The "Cambrian substrate revolution" or "Agronomic revolution", evidenced in trace fossils, is the diversification of animal burrowing during the early Cambrian period. Before this "widening of the behavioural repertoire", bottom-dwelling anima ...
), implying that sufficiently complex organisms were common over 1 billion years ago. The connection between grazer and stromatolite abundance is well documented in the younger
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 ...
evolutionary radiation An evolutionary radiation is an increase in taxonomic diversity that is caused by elevated rates of speciation, that may or may not be associated with an increase in morphological disparity. Radiations may affect one clade or many, and be rapid ...
; stromatolite abundance also increased after the end-Ordovician and end-Permian extinctions decimated marine animals, falling back to earlier levels as marine animals recovered. Fluctuations in
metazoan Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...
population and diversity may not have been the only factor in the reduction in stromatolite abundance. Factors such as the chemistry of the environment may have been responsible for changes. While
prokaryotic A prokaryote () is a single-celled organism that lacks a nucleus and other membrane-bound organelles. The word ''prokaryote'' comes from the Greek πρό (, 'before') and κάρυον (, 'nut' or 'kernel').Campbell, N. "Biology:Concepts & Connec ...
cyanobacteria themselves reproduce asexually through cell division, they were instrumental in priming the environment for the evolutionary development of more complex
eukaryotic Eukaryotes () are organisms whose Cell (biology), cells have a cell nucleus, 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 ...
organisms. Cyanobacteria (as well as
extremophile An extremophile (from Latin ' meaning "extreme" and Greek ' () meaning "love") is an organism that is able to live (or in some cases thrive) in extreme environments, i.e. environments that make survival challenging such as due to extreme temper ...
Gammaproteobacteria) are thought to be largely responsible for increasing the amount of
oxygen Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements as ...
in the primeval earth's atmosphere through their continuing
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 i ...
. Cyanobacteria use
water Water (chemical formula ) is an Inorganic compound, inorganic, transparent, tasteless, odorless, and Color of water, nearly colorless chemical substance, which is the main constituent of Earth's hydrosphere and the fluids of all known living ...
,
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 trans ...
and sunlight to create their food. A layer of
mucus Mucus ( ) is a slippery aqueous secretion produced by, and covering, mucous membranes. It is typically produced from cells found in mucous glands, although it may also originate from mixed glands, which contain both serous and mucous cells. It ...
often forms over mats of cyanobacterial cells. In modern microbial mats, debris from the surrounding habitat can become trapped within the mucus, which can be cemented by the calcium carbonate to grow thin laminations of
limestone Limestone ( calcium carbonate ) is a type of carbonate sedimentary rock which is the main source of the material lime. It is composed mostly of the minerals calcite and aragonite, which are different crystal forms of . Limestone forms whe ...
. These laminations can accrete over time, resulting in the banded pattern common to stromatolites. The domal morphology of biological stromatolites is the result of the vertical growth necessary for the continued infiltration of sunlight to the organisms for photosynthesis. Layered spherical growth structures termed
oncolite Oncolites are sedimentary structures composed of oncoids, which are layered structures formed by cyanobacterial growth. Oncolites are very similar to stromatolites, but, instead of forming columns, they form approximately spherical structures. T ...
s are similar to stromatolites and are also known from the
fossil record 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 ...
.
Thrombolite Thrombolites (from Ancient Greek θρόμβος ''thrómbos'' meaning " clot" and λῐ́θος ''líthos'' meaning "stone") are clotted accretionary structures formed in shallow water by the trapping, binding, and cementation of sedimentary g ...
s are poorly laminated or non-laminated clotted structures formed by cyanobacteria common in the fossil record and in modern sediments. The Zebra River Canyon area of the Kubis platform in the deeply dissected Zaris Mountains of southwestern
Namibia Namibia (, ), officially the Republic of Namibia, is a country in Southern Africa. Its western border is the Atlantic Ocean. It shares land borders with Zambia and Angola to the north, Botswana to the east and South Africa to the south and ea ...
provides an extremely well exposed example of the thrombolite-stromatolite-metazoan reefs that developed during the Proterozoic period, the stromatolites here being better developed in updip locations under conditions of higher current velocities and greater sediment influx.


Astrobiology

It has been suggested that biominerals could be important indicators of extraterrestrial life and thus could play an important role in the search for past or present life on the planet
Mars Mars is the fourth planet from the Sun and the second-smallest planet in the Solar System, only being larger than Mercury. In the English language, Mars is named for the Roman god of war. Mars is a terrestrial planet with a thin at ...
. Furthermore, organic components ( biosignatures) that are often associated with biominerals are believed to play crucial roles in both pre-biotic and biotic reactions. On 24 January 2014, NASA reported that current studies by the ''Curiosity'' and ''Opportunity'' rovers on Mars will now be searching for evidence of ancient life, including a
biosphere The biosphere (from Greek βίος ''bíos'' "life" and σφαῖρα ''sphaira'' "sphere"), also known as the ecosphere (from Greek οἶκος ''oîkos'' "environment" and σφαῖρα), is the worldwide sum of all ecosystems. It can also ...
based on
autotroph An autotroph or primary producer is an organism that produces complex organic compounds (such as carbohydrates, fats, and proteins) using carbon from simple substances such as carbon dioxide,Morris, J. et al. (2019). "Biology: How Life Wo ...
ic,
chemotroph A Chemotroph is an organism that obtains energy by the oxidation of electron donors in their environments. These molecules can be organic (chemoorganotrophs) or inorganic ( chemolithotrophs). The chemotroph designation is in contrast to phototr ...
ic and/or
chemolithoautotrophic A lithoautotroph is an organism which derives energy from reactions of reduced compounds of mineral (inorganic) origin. Two types of lithoautotrophs are distinguished by their energy source; photolithoautotrophs derive their energy from light while ...
microorganism A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in olde ...
s, as well as ancient water, including fluvio-lacustrine environments (
plain In geography, a plain is a flat expanse of land that generally does not change much in elevation, and is primarily treeless. Plains occur as lowlands along valleys or at the base of mountains, as coastal plains, and as plateaus or uplands ...
s related to ancient
river A river is a natural flowing watercourse, usually freshwater, flowing towards an ocean, sea, lake or another river. In some cases, a river flows into the ground and becomes dry at the end of its course without reaching another body of w ...
s or
lake A lake is an area filled with water, localized in a basin, surrounded by land, and distinct from any river or other outlet that serves to feed or drain the lake. Lakes lie on land and are not part of the ocean, although, like the much large ...
s) that may have been
habitable Habitability refers to the adequacy of an environment for human living. Where housing is concerned, there are generally local ordinances which define habitability. If a residence complies with those laws it is said to be habitable. In extreme e ...
. The search for evidence of
habitability Habitability refers to the adequacy of an environment for human living. Where housing is concerned, there are generally local ordinances which define habitability. If a residence complies with those laws it is said to be habitable. In extreme e ...
,
taphonomy Taphonomy is the study of how organisms decay and become fossilized or preserved in the paleontological record. The term ''taphonomy'' (from Greek , 'burial' and , 'law') was introduced to paleontology in 1940 by Soviet scientist Ivan Efremov t ...
(related to fossils), and
organic carbon Total organic carbon (TOC) is the amount of carbon found in an organic compound and is often used as a non-specific indicator of water quality or cleanliness of pharmaceutical manufacturing equipment. TOC may also refer to the amount of organic c ...
on the planet
Mars Mars is the fourth planet from the Sun and the second-smallest planet in the Solar System, only being larger than Mercury. In the English language, Mars is named for the Roman god of war. Mars is a terrestrial planet with a thin at ...
is now a primary
NASA The National Aeronautics and Space Administration (NASA ) is an independent agencies of the United States government, independent agency of the US federal government responsible for the civil List of government space agencies, space program ...
objective.


Pseudofossils

''Pseudofossils'' are visual patterns in rocks that are produced by geologic processes rather than biologic processes. They can easily be mistaken for real fossils. Some pseudofossils, such as geological
dendrite Dendrites (from Greek δένδρον ''déndron'', "tree"), also dendrons, are branched protoplasmic extensions of a nerve cell that propagate the electrochemical stimulation received from other neural cells to the cell body, or soma, of the ...
crystals, are formed by naturally occurring fissures in the rock that get filled up by percolating minerals. Other types of pseudofossils are kidney ore (round shapes in iron ore) and moss agates, which look like moss or plant leaves.
Concretion A concretion is a hard, compact mass of matter formed by the precipitation of mineral cement within the spaces between particles, and is found in sedimentary rock or soil. Concretions are often ovoid or spherical in shape, although irregular ...
s, spherical or ovoid-shaped nodules found in some sedimentary strata, were once thought to be
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
eggs, and are often mistaken for fossils as well.


History of the study of fossils

Gathering fossils dates at least to the beginning of recorded history. The fossils themselves are referred to as the fossil record. The fossil record was one of the early sources of data underlying the study of
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 ...
and continues to be relevant to the history of life on Earth.
Paleontologist Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
s examine the fossil record to understand the process of evolution and the way particular
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
have evolved.


Ancient civilizations

Fossils have been visible and common throughout most of natural history, and so documented human interaction with them goes back as far as recorded history, or earlier. There are many examples of paleolithic stone knives in Europe, with fossil
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 ...
s set precisely at the hand grip, going all the way back to ''
Homo heidelbergensis ''Homo heidelbergensis'' (also ''H. sapiens heidelbergensis''), sometimes called Heidelbergs, is an extinct species or subspecies of archaic human which existed during the Middle Pleistocene. It was subsumed as a subspecies of '' H. erectus'' i ...
'' and
Neanderthal Neanderthals (, also ''Homo neanderthalensis'' and erroneously ''Homo sapiens neanderthalensis''), also written as Neandertals, are an extinct species or subspecies of archaic humans who lived in Eurasia until about 40,000 years ago. While the ...
s. These ancient peoples also drilled holes through the center of those round fossil shells, apparently using them as beads for necklaces. The ancient Egyptians gathered fossils of species that resembled the bones of modern species they worshipped. The god
Set Set, The Set, SET or SETS may refer to: Science, technology, and mathematics Mathematics *Set (mathematics), a collection of elements *Category of sets, the category whose objects and morphisms are sets and total functions, respectively Electro ...
was associated with the
hippopotamus The hippopotamus ( ; : hippopotamuses or hippopotami; ''Hippopotamus amphibius''), also called the hippo, common hippopotamus, or river hippopotamus, is a large semiaquatic mammal native to sub-Saharan Africa. It is one of only two extan ...
, therefore fossilized bones of hippo-like species were kept in that deity's temples. Five-rayed fossil sea urchin shells were associated with the deity
Sopdu Sopdu (also rendered Septu or Sopedu) was a god of the sky and of eastern border regions in the religion of Ancient Egypt.Wilkinson, Richard H. (2003). ''The Complete Gods and Goddesses of Ancient Egypt''. Thames & Hudson. p. 211 He was Khensit's ...
, the Morning Star, equivalent of
Venus Venus is the second planet from the Sun. It is sometimes called Earth's "sister" or "twin" planet as it is almost as large and has a similar composition. As an interior planet to Earth, Venus (like Mercury) appears in Earth's sky never f ...
in Roman mythology. Fossils appear to have directly contributed to the mythology of many civilizations, including the ancient Greeks. Classical Greek historian Herodotos wrote of an area near
Hyperborea In Greek mythology, the Hyperboreans ( grc, Ὑπερβόρε(ι)οι, ; la, Hyperborei) were a mythical people who lived in the far northern part of the known world. Their name appears to derive from the Greek , "beyond Boreas" (the God of ...
where
gryphon The griffin, griffon, or gryphon ( Ancient Greek: , ''gryps''; Classical Latin: ''grȳps'' or ''grȳpus''; Late and Medieval Latin: ''gryphes'', ''grypho'' etc.; Old French: ''griffon'') is a legendary creature with the body, tail, and ...
s protected golden treasure. There was indeed gold mining in that approximate region, where beaked ''
Protoceratops ''Protoceratops'' (; ) is a genus of small protoceratopsid dinosaurs that lived in Asia during the Late Cretaceous, around 75 to 71 million years ago. The genus ''Protoceratops'' includes two species: ''P. andrewsi'' and the larger ''P. hellenik ...
'' skulls were common as fossils. A later
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
scholar,
Aristotle Aristotle (; grc-gre, Ἀριστοτέλης ''Aristotélēs'', ; 384–322 BC) was a Greek philosopher and polymath during the Classical period in Ancient Greece. Taught by Plato, he was the founder of the Peripatetic school of ph ...
, eventually realized that fossil seashells from rocks were similar to those found on the beach, indicating the fossils were once living animals. He had previously explained them in terms of
vapor In physics, a vapor (American English) or vapour (British English and Canadian English; see spelling differences) is a substance in the gas phase at a temperature lower than its critical temperature,R. H. Petrucci, W. S. Harwood, and F. G. Her ...
ous
exhalation Exhalation (or expiration) is the flow of the breath out of an organism. In animals, it is the movement of air from the lungs out of the airways, to the external environment during breathing. This happens due to elastic properties of the lungs, ...
s, which
Persia Iran, officially the Islamic Republic of Iran, and also called Persia, is a country located in Western Asia. It is bordered by Iraq and Turkey to the west, by Azerbaijan and Armenia to the northwest, by the Caspian Sea and Turkmeni ...
n polymath Avicenna modified into the theory of petrifying fluids (). Recognition of fossil seashells as originating in the sea was built upon in the 14th century by
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 ...
, and accepted in some form by most naturalists by the 16th century. Roman naturalist
Pliny the Elder Gaius Plinius Secundus (AD 23/2479), called Pliny the Elder (), was a Roman author, naturalist and natural philosopher, and naval and army commander of the early Roman Empire, and a friend of the emperor Vespasian. He wrote the encyclopedic ' ...
wrote of " tongue stones", which he called glossopetra. These were fossil shark teeth, thought by some classical cultures to look like the tongues of people or snakes. He also wrote about the
horns of Ammon The horns of Ammon were curling ram horns, used as a symbol of the Egyptian deity Ammon (also spelled Amun or Amon). Because of the visual similarity, they were also associated with the fossils shells of ancient snails and cephalopods, the latter ...
, which are fossil ammonites, from whence the group of shelled octopus-cousins ultimately draws its modern name. Pliny also makes one of the earlier known references to
toadstone The toadstone, also known as bufonite (from Latin , "toad"), is a mythical stone or gem that was thought to be found in the head of a toad. It was supposed to be an antidote to poison and in this it is like batrachite, supposedly formed in the ...
s, thought until the 18th century to be a magical cure for poison originating in the heads of toads, but which are fossil teeth from ''
Lepidotes ''Lepidotes'' (from el, λεπιδωτός , 'covered with scales') (previously known as ''Lepidotus'') is an extinct genus of Mesozoic ray-finned fish. It has been considered a wastebasket taxon, characterised by "general features, such as thi ...
'', a
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 ...
ray-finned fish. The
Plains tribes Plains Indians or Indigenous peoples of the Great Plains and Canadian Prairies are the Native American tribes and First Nation band governments who have historically lived on the Interior Plains (the Great Plains and Canadian Prairies) ...
of North America are thought to have similarly associated fossils, such as the many intact pterosaur fossils naturally exposed in the region, with their own mythology of the
thunderbird Thunderbird, thunder bird or thunderbirds may refer to: * Thunderbird (mythology), a legendary creature in certain North American indigenous peoples' history and culture * Ford Thunderbird, a car Birds * Dromornithidae, extinct flightless birds ...
. There is no such direct mythological connection known from prehistoric Africa, but there is considerable evidence of tribes there excavating and moving fossils to ceremonial sites, apparently treating them with some reverence. In Japan, fossil shark teeth were associated with the mythical
tengu are a type of legendary creature found in Japanese folk religion (Shinto). They are considered a type of '' yōkai'' (supernatural beings) or Shinto ''kami'' (gods). The ''Tengu'' were originally thought to take the forms of birds of prey and ...
, thought to be the razor-sharp claws of the creature, documented some time after the 8th century AD. In medieval China, the fossil bones of ancient mammals including '' Homo erectus'' were often mistaken for " dragon bones" and used as medicine and
aphrodisiac An aphrodisiac is a substance that increases sexual desire, sexual attraction, sexual pleasure, or sexual behavior. Substances range from a variety of plants, spices, foods, and synthetic chemicals. Natural aphrodisiacs like cannabis or cocai ...
s. In addition, some of these fossil bones are collected as "art" by scholars, who left scripts on various artifacts, indicating the time they were added to a collection. One good example is the famous scholar
Huang Tingjian Huang Tingjian (; 1045–1105) was a Chinese calligrapher, painter, and poet of the Song dynasty. He is predominantly known as a calligrapher, and is also admired for his painting and poetry. He was one of the Four Masters of the Song Dynasty () ...
of the
Song Dynasty The Song dynasty (; ; 960–1279) was an imperial dynasty of China that began in 960 and lasted until 1279. The dynasty was founded by Emperor Taizu of Song following his usurpation of the throne of the Later Zhou. The Song conquered the rest ...
during the 11th century, who kept a specific seashell fossil with his own poem engraved on it. In his '' Dream Pool Essays'' published in 1088, Song dynasty Chinese
scholar-official The scholar-officials, also known as literati, scholar-gentlemen or scholar-bureaucrats (), were government officials and prestigious scholars in Chinese society, forming a distinct social class. Scholar-officials were politicians and governmen ...
Shen Kuo hypothesized that marine fossils found in a geological stratum of mountains located hundreds of miles from the
Pacific Ocean The Pacific Ocean is the largest and deepest of Earth's five oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean (or, depending on definition, to Antarctica) in the south, and is bounded by the contin ...
was evidence that a prehistoric seashore had once existed there and shifted over centuries of time.Needham, Joseph. (1959). ''Science and Civilization in China: Volume 3, Mathematics and the Sciences of the Heavens and the Earth''.
Cambridge University Press Cambridge University Press is the university press of the University of Cambridge. Granted letters patent by King Henry VIII in 1534, it is the oldest university press in the world. It is also the King's Printer. Cambridge University Pre ...
. pp. 603–618.
His observation 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 proce ...
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, ...
s in the dry northern climate zone of what is now
Yan'an Yan'an (; ), alternatively spelled as Yenan is a prefecture-level city in the Shaanbei region of Shaanxi province, China, bordering Shanxi to the east and Gansu to the west. It administers several counties, including Zhidan (formerly Bao'an) ...
,
Shaanxi Shaanxi (alternatively Shensi, see § Name) is a landlocked province of China. Officially part of Northwest China, it borders the province-level divisions of Shanxi (NE, E), Henan (E), Hubei (SE), Chongqing (S), Sichuan (SW), Gansu (W), N ...
province, China, led him to advance early ideas of gradual
climate change In common usage, climate change describes global warming—the ongoing increase in global average temperature—and its effects on Earth's climate system. Climate change in a broader sense also includes previous long-term changes to ...
due to bamboo naturally growing in wetter climate areas.Rafferty, John P. (2012). ''Geological Sciences; Geology: Landforms, Minerals, and Rocks''. New York: Britannica Educational Publishing, p. 6. In medieval
Christendom Christendom historically refers to the Christian states, Christian-majority countries and the countries in which Christianity dominates, prevails,SeMerriam-Webster.com : dictionary, "Christendom"/ref> or is culturally or historically intertwine ...
, fossilized sea creatures on mountainsides were seen as proof of the biblical deluge of
Noah's Ark Noah's Ark ( he, תיבת נח; Biblical Hebrew: ''Tevat Noaḥ'')The word "ark" in modern English comes from Old English ''aerca'', meaning a chest or box. (See Cresswell 2010, p.22) The Hebrew word for the vessel, ''teva'', occurs twice in ...
. After observing the existence of seashells in mountains, the
ancient Greek philosopher Ancient Greek philosophy arose in the 6th century BC, marking the end of the Greek Dark Ages. Greek philosophy continued throughout the Hellenistic period and the period in which Greece and most Greek-inhabited lands were part of the Roman Empire ...
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 Classical ...
(c. 570 – 478 BC) speculated that the world was once inundated in a great flood that buried living creatures in drying mud.Rafferty, John P. (2012). ''Geological Sciences; Geology: Landforms, Minerals, and Rocks''. New York: Britannica Educational Publishing, pp. 5–6. . In 1027, the
Persian Persian may refer to: * People and things from Iran, historically called ''Persia'' in the English language ** Persians, the majority ethnic group in Iran, not to be conflated with the Iranic peoples ** Persian language, an Iranian language of the ...
Avicenna explained fossils' stoniness in ''
The Book of Healing ''The Book of Healing'' (; ; also known as ) is a scientific and philosophical encyclopedia written by Abu Ali ibn Sīna (aka Avicenna) from medieval Persia, near Bukhara in Maverounnahr. He most likely began to compose the book in 1014, comp ...
'': From the 13th century to the present day, scholars pointed out that the fossil skulls of Deinotherium giganteum, found in
Crete Crete ( el, Κρήτη, translit=, Modern: , Ancient: ) is the largest and most populous of the Greek islands, the 88th largest island in the world and the fifth largest island in the Mediterranean Sea, after Sicily, Sardinia, Cyprus, ...
and Greece, might have been interpreted as being the skulls of the
Cyclopes In Greek mythology and later Roman mythology, the Cyclopes ( ; el, Κύκλωπες, ''Kýklōpes'', "Circle-eyes" or "Round-eyes"; singular Cyclops ; , ''Kýklōps'') are giant one-eyed creatures. Three groups of Cyclopes can be distinguish ...
of
Greek mythology A major branch of classical mythology, Greek mythology is the body of myths originally told by the ancient Greeks, and a genre of Ancient Greek folklore. These stories concern the origin and nature of the world, the lives and activities ...
, and are possibly the origin of that Greek myth. Their skulls appear to have a single eye-hole in the front, just like their modern
elephant Elephants are the largest existing land animals. Three living species are currently recognised: the African bush elephant, the African forest elephant, and the Asian elephant. They are the only surviving members of the family Elephantidae ...
cousins, though in fact it's actually the opening for their trunk. In Norse mythology, echinoderm shells (the round five-part button left over from a sea urchin) were associated with the god
Thor Thor (; from non, Þórr ) is a prominent god in Germanic paganism. In Norse mythology, he is a hammer-wielding god associated with lightning, thunder, storms, sacred groves and trees, strength, the protection of humankind, hallowing, an ...
, not only being incorporated in thunderstones, representations of Thor's hammer and subsequent hammer-shaped crosses as Christianity was adopted, but also kept in houses to garner Thor's protection. These grew into the shepherd's crowns of English folklore, used for decoration and as good luck charms, placed by the doorway of homes and churches. In Suffolk, a different species was used as a good-luck charm by bakers, who referred to them as fairy loaves, associating them with the similarly shaped loaves of bread they baked.


Early modern explanations

More scientific views of fossils emerged during the
Renaissance The Renaissance ( , ) , from , with the same meanings. is a period in European history The history of Europe is traditionally divided into four time periods: prehistoric Europe (prior to about 800 BC), classical antiquity (800 BC to AD ...
.
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, Drawing, draughtsman, engineer, scientist, theorist, sculptor, and architect. While his fame initially res ...
concurred with Aristotle's view that fossils were the remains of ancient life. For example, Leonardo noticed discrepancies with the biblical flood narrative as an explanation for fossil origins: In 1666,
Nicholas Steno Niels Steensen ( da, Niels Steensen; Latinized to ''Nicolaus Steno'' or ''Nicolaus Stenonius''; 1 January 1638 – 25 November 1686 Robert Hooke (1635–1703) included
micrograph A micrograph or photomicrograph is a photograph or digital image taken through a microscope or similar device to show a magnified image of an object. This is opposed to a macrograph or photomacrograph, an image which is also taken on a m ...
s of fossils in his ''
Micrographia ''Micrographia: or Some Physiological Descriptions of Minute Bodies Made by Magnifying Glasses. With Observations and Inquiries Thereupon.'' is a historically significant book by Robert Hooke about his observations through various lenses. It ...
'' and was among the first to observe fossil
foram Foraminifera (; Latin for "hole bearers"; informally called "forams") are single-celled organisms, members of a phylum or class of amoeboid protists characterized by streaming granular ectoplasm for catching food and other uses; and commonly a ...
s. His observations on fossils, which he stated to be the petrified remains of creatures some of which no longer existed, were published posthumously in 1705. William Smith (1769–1839), an English canal engineer, observed that rocks of different ages (based on the
law of superposition The law of superposition is an axiom that forms one of the bases of the sciences of geology, archaeology, and other fields pertaining to geological stratigraphy. In its plainest form, it states that in undeformed stratigraphic sequences, the ...
) preserved different assemblages of fossils, and that these assemblages succeeded one another in a regular and determinable order. He observed that rocks from distant locations could be correlated based on the fossils they contained. He termed this the principle of ''faunal succession''. This principle became one of Darwin's chief pieces of evidence that biological evolution was real. Georges Cuvier came to believe that most if not all the animal fossils he examined were remains of extinct species. This led Cuvier to become an active proponent of the geological school of thought called catastrophism. Near the end of his 1796 paper on living and fossil elephants he said: Interest in fossils, and geology more generally, expanded during the early nineteenth century. In Britain,
Mary Anning Mary Anning (21 May 1799 – 9 March 1847) was an English fossil collector, dealer, and palaeontologist who became known around the world for the discoveries she made in Jurassic marine fossil beds in the cliffs along the English Channel ...
's discoveries of fossils, including the first complete ichthyosaur and a complete
plesiosaurus ''Plesiosaurus'' (Greek: ' ('), near to + ' ('), lizard) is a genus of extinct, large marine sauropterygian reptile that lived during the Early Jurassic. It is known by nearly complete skeletons from the Lias of England. It is distinguishable b ...
skeleton, sparked both public and scholarly interest.


Linnaeus and Darwin

Early naturalists well understood the similarities and differences of living species leading
Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, the ...
to develop a hierarchical classification system still in use today. Darwin and his contemporaries first linked the hierarchical structure of the tree of life with the then very sparse fossil record. Darwin eloquently described a process of descent with modification, or evolution, whereby organisms either adapt to natural and changing environmental pressures, or they perish. When Darwin wrote '' On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life'', the oldest animal fossils were those from the Cambrian Period, now known to be about 540 million years old. He worried about the absence of older fossils because of the implications on the validity of his theories, but he expressed hope that such fossils would be found, noting that: "only a small portion of the world is known with accuracy." Darwin also pondered the sudden appearance of many groups (i.e. phyla) in the oldest known Cambrian fossiliferous strata.


After Darwin

Since Darwin's time, the fossil record has been extended to between 2.3 and 3.5 billion years. Most of these Precambrian fossils are microscopic bacteria or
microfossils A microfossil is a fossil that is generally between 0.001 mm and 1 mm in size, the visual study of which requires the use of light or electron microscopy. A fossil which can be studied with the naked eye or low-powered magnification, ...
. However, macroscopic fossils are now known from the late Proterozoic. The
Ediacara 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 sessil ...
(also called Vendian biota) dating from 575 million years ago collectively constitutes a richly diverse assembly of early multicellular eukaryotes. The fossil record and faunal succession form the basis of the science 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 B ...
or determining the age of rocks based on embedded fossils. For the first 150 years of
geology Geology () is a branch of natural science concerned with Earth and other astronomical objects, the features or rocks of which it is composed, and the processes by which they change over time. Modern geology significantly overlaps all other Ea ...
, biostratigraphy and superposition were the only means for determining the relative age of rocks. 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 ...
was developed based on the relative ages of rock strata as determined by the early paleontologists and stratigraphers. Since the early years of the twentieth century,
absolute dating Absolute dating is the process of determining an age on a specified chronology in archaeology and geology. Some scientists prefer the terms chronometric or calendar dating, as use of the word "absolute" implies an unwarranted certainty of accuracy ...
methods, such as
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 ...
(including potassium/argon, argon/argon,
uranium series In nuclear science, the decay chain refers to a series of radioactive decays of different radioactive decay products as a sequential series of transformations. It is also known as a "radioactive cascade". Most radioisotopes do not decay direct ...
, and, for very recent fossils, radiocarbon dating) have been used to verify the relative ages obtained by fossils and to provide absolute ages for many fossils. Radiometric dating has shown that the earliest known stromatolites are over 3.4 billion years old.


Modern era

Paleontology has joined with
evolutionary biology Evolutionary biology is the subfield of biology that studies the evolutionary processes ( natural selection, common descent, speciation) that produced the diversity of life on Earth. It is also defined as the study of the history of life ...
to share the interdisciplinary task of outlining the tree of life, which inevitably leads backwards in time to Precambrian microscopic life when cell structure and functions evolved. Earth's deep time in the Proterozoic and deeper still in the Archean is only "recounted by microscopic fossils and subtle chemical signals." Molecular biologists, using
phylogenetics In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups ...
, can compare protein
amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha a ...
or
nucleotide Nucleotides are organic molecules consisting of a nucleoside and a phosphate. They serve as monomeric units of the nucleic acid polymers – deoxyribonucleic acid (DNA) and ribonucleic acid (RNA), both of which are essential biomolecule ...
sequence homology (i.e., similarity) to evaluate taxonomy and evolutionary distances among organisms, with limited statistical confidence. The study of fossils, on the other hand, can more specifically pinpoint when and in what organism a mutation first appeared. Phylogenetics and paleontology work together in the clarification of science's still dim view of the appearance of life and its evolution.
Niles Eldredge Niles Eldredge (; born August 25, 1943) is an American biologist and paleontologist, who, along with Stephen Jay Gould, proposed the theory of punctuated equilibrium in 1972. Education Eldredge began his undergraduate studies in Latin at Columb ...
's study of the ''
Phacops ''Phacops'' is a genus of trilobites in the order Phacopida, family Phacopidae, that lived in Europe, northwestern Africa, North and South America and China from the Late Ordovician until the very end of the Devonian, with a broader time range ...
''
trilobite Trilobites (; meaning "three lobes") are extinct marine arthropods that form the class Trilobita. Trilobites form one of the earliest-known groups of arthropods. The first appearance of trilobites in the fossil record defines the base of the ...
genus supported the hypothesis that modifications to the arrangement of the trilobite's eye lenses proceeded by fits and starts over millions of years during the Devonian. Eldredge's interpretation of the ''Phacops'' fossil record was that the aftermaths of the lens changes, but not the rapidly occurring evolutionary process, were fossilized. This and other data led Stephen Jay Gould and Niles Eldredge to publish their seminal paper on
punctuated equilibrium In evolutionary biology, punctuated equilibrium (also called punctuated equilibria) is a theory that proposes that once a species appears in the fossil record, the population will become stable, showing little evolutionary change for most of i ...
in 1971. Synchrotron
X-ray An X-ray, or, much less commonly, X-radiation, is a penetrating form of high-energy electromagnetic radiation. Most X-rays have a wavelength ranging from 10  picometers to 10  nanometers, corresponding to frequencies in the range 30&nb ...
tomographic analysis of early Cambrian bilaterian embryonic microfossils yielded new insights of
metazoan Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...
evolution at its earliest stages. The tomography technique provides previously unattainable three-dimensional resolution at the limits of fossilization. Fossils of two enigmatic bilaterians, the worm-like ''
Markuelia ''Markuelia'' is a genus of fossil worm-like bilaterian animals allied to Ecdysozoa and known from strata of Lower Cambrian to Lower Ordovician age containing five species. An advanced X-ray imaging technique called X-ray tomographic microscop ...
'' and a putative, primitive
protostome Protostomia () is the clade of animals once thought to be characterized by the formation of the organism's mouth before its anus during embryogenesis, embryonic development. This nature has since been discovered to be extremely variable among Pro ...
, '' Pseudooides'', provide a peek at germ layer embryonic development. These 543-million-year-old embryos support the emergence of some aspects of
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 ...
development earlier than previously thought in the late Proterozoic. The preserved embryos from China and
Siberia Siberia ( ; rus, Сибирь, r=Sibir', p=sʲɪˈbʲirʲ, a=Ru-Сибирь.ogg) is an extensive region, geographical region, constituting all of North Asia, from the Ural Mountains in the west to the Pacific Ocean in the east. It has been a ...
underwent rapid
diagenetic Diagenesis () is the process that describes physical and chemical changes in sediments first caused by water-rock interactions, microbial activity, and compaction after their deposition. Increased pressure and temperature only start to play a ...
phosphatization resulting in exquisite preservation, including cell structures. This research is a notable example of how knowledge encoded by the fossil record continues to contribute otherwise unattainable information on the emergence and development of life on Earth. For example, the research suggests ''Markuelia'' has closest affinity to priapulid worms, and is adjacent to the evolutionary branching of
Priapulida Priapulida (priapulid worms, from Gr. πριάπος, ''priāpos'' 'Priapus' + Lat. ''-ul-'', diminutive), sometimes referred to as penis worms, is a phylum of unsegmented marine worms. The name of the phylum relates to the Greek god of fertility ...
,
Nematoda The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-parasitic nematodes also known as eelworms. They are a diverse animal phylum inhabiting a broa ...
and
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 ...
a. Despite significant advances in uncovering and identifying paleontological specimens, it is generally accepted that the fossil record is vastly incomplete. Approaches for measuring the completeness of the fossil record have been developed for numerous subsets of species, including those grouped taxonomically, temporally, environmentally/geographically, or in sum. This encompasses the subfield of
taphonomy Taphonomy is the study of how organisms decay and become fossilized or preserved in the paleontological record. The term ''taphonomy'' (from Greek , 'burial' and , 'law') was introduced to paleontology in 1940 by Soviet scientist Ivan Efremov t ...
and the study of biases in the paleontological record.


Art

According to one hypothesis, a Corinthian vase from the 6th century B.C. C. is the oldest artistic record of a vertebrate fossil, perhaps a Miocene giraffe combined with elements from other species. However, a subsequent study using artificial intelligence and expert evaluations reject this idea, because mammals do not have the eye bones shown in the painted monster. Morphologically, the vase painting correspond to a carnivorous reptile of the Varanidae family that still lives in regions occupied by the ancient Greek.


Trading and collecting

Fossil trading is the practice of buying and selling fossils. This is many times done illegally with artifacts stolen from research sites, costing many important scientific specimens each year. The problem is quite pronounced in China, where many specimens have been stolen. Fossil collecting (sometimes, in a non-scientific sense, fossil hunting) is the collection of fossils for scientific study, hobby, or profit. Fossil collecting, as practiced by amateurs, is the predecessor of modern paleontology and many still collect fossils and study fossils as amateurs. Professionals and amateurs alike collect fossils for their scientific value.


As medicine

The use of fossils to address health issues is rooted in
traditional medicine Traditional medicine (also known as indigenous medicine or folk medicine) comprises medical aspects of traditional knowledge that developed over generations within the folk beliefs of various societies, including indigenous peoples, before the ...
and include the use of fossils as
talisman A talisman is any object ascribed with religious or magical powers intended to protect, heal, or harm individuals for whom they are made. Talismans are often portable objects carried on someone in a variety of ways, but can also be installed perm ...
s. The specific fossil to use to alleviate or cure an illness is often based on its resemblance to the symptoms or affected organ. The usefulness of fossils as medicine is almost entirely a
placebo effect A placebo ( ) is a substance or treatment which is designed to have no therapeutic value. Common placebos include inert tablets (like sugar pills), inert injections (like Saline (medicine), saline), sham surgery, and other procedures. In general ...
, though fossil material might conceivably have some
antacid An antacid is a substance which neutralizes stomach acidity and is used to relieve heartburn, indigestion or an upset stomach. Some antacids have been used in the treatment of constipation and diarrhea. Marketed antacids contain salts of alu ...
activity or supply some essential minerals. The use of dinosaur bones as "dragon bones" has persisted in
Traditional Chinese medicine Traditional Chinese medicine (TCM) is an alternative medical practice drawn from traditional medicine in China. It has been described as "fraught with pseudoscience", with the majority of its treatments having no logical mechanism of acti ...
into modern times, with mid-Cretaceous dinosaur bones being used for the purpose in
Ruyang County Ruyang County () is a county in the west of Henan province, China, under the jurisdiction of the prefecture-level city of Luoyang. Ruyang County has been the site of several major discoveries of fossilized dinosaur bones, including ''Huanghetit ...
during the early 21st century.


Gallery

File:Marine fossils found high in the Himalayas. Collection of the Abbot of Dhankar Gompa, HP, India.jpg, Marine fossils found high in the Himalayas. Collection of the Abbot of
Dhankar Gompa Dhankar Gompa (also Dankhar, Drangkhar or Dhangkar Gompa; ''Brang-mkhar'' or ''Grang-mkhar'') is a village and also a Gompa, a Buddhist temple in the district of Lahaul and Spiti in India. It is situated at an elevation of 3,894 metres (12,7 ...
, HP, India File:Amonite Cropped.jpg, Three small ammonite fossils, each approximately 1.5 cm across File:Priscacara liops Green River Formation.jpg,
Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', " ...
fossil fish ''Priscacara liops'' from the
Green River Formation The Green River Formation is an Eocene geologic formation that records the sedimentation in a group of intermountain lakes in three basins along the present-day Green River in Colorado, Wyoming, and Utah. The sediments are deposited in very fine ...
of Wyoming File:Trilobite2.jpg, A permineralized
trilobite Trilobites (; meaning "three lobes") are extinct marine arthropods that form the class Trilobita. Trilobites form one of the earliest-known groups of arthropods. The first appearance of trilobites in the fossil record defines the base of the ...
, ''
Asaphus kowalewskii ''Asaphus kowalewskii'' () is one of the 35 species of trilobites of the genus ''Asaphus'' (this particular species is sometimes placed in its own genus, ''Neoasaphus''). Fossils of this species are popular among collectors because of their promi ...
'' File:Carcharodontosaurus and Megalodon teeth.jpg,
Megalodon Megalodon (''Otodus megalodon''), meaning "big tooth", is an extinct species of mackerel shark that lived approximately 23 to 3.6 million years ago (Mya), from the Early Miocene to the Pliocene epochs. It was formerly thought to be a membe ...
and ''
Carcharodontosaurus ''Carcharodontosaurus'' (; ) is a genus of large carcharodontosaurid theropod dinosaur that existed during the Cenomanian age of the Late Cretaceous in Northern Africa. The genus ''Carcharodontosaurus'' is named after the shark genus '' Carc ...
'' teeth. The latter was found in the
Sahara Desert , photo = Sahara real color.jpg , photo_caption = The Sahara taken by Apollo 17 astronauts, 1972 , map = , map_image = , location = , country = , country1 = , ...
. File:The fossils from Cretaceous age found in Lebanon.jpg, Fossil shrimp (
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 ...
) File:PetrifiedWood.jpg, Petrified wood in Petrified Forest National Park, Arizona File:Petrified Araucaria cone from patagonia-Edit3.jpg, Petrified
cone A cone is a three-dimensional geometric shape that tapers smoothly from a flat base (frequently, though not necessarily, circular) to a point called the apex or vertex. A cone is formed by a set of line segments, half-lines, or lines con ...
of ''
Araucaria mirabilis ''Araucaria mirabilis'' is an extinct species of coniferous tree from Patagonia, Argentina. It belongs to the genus '' Araucaria''. ''A. mirabilis'' are known from large amounts of very well preserved silicified wood and cones from the Cerro ...
'' from
Patagonia Patagonia () refers to a geographical region that encompasses the southern end of South America, governed by Argentina and Chile. The region comprises the southern section of the Andes Mountains with lakes, fjords, temperate rainforests, and g ...
,
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
dating from the
Jurassic Period The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
(approx. 210 Ma) File:CyprusPlioceneGastropod.JPG, A fossil gastropod from the
Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58Cyprus Cyprus ; tr, Kıbrıs (), officially the Republic of Cyprus,, , lit: Republic of Cyprus is an island country located south of the Anatolian Peninsula in the eastern Mediterranean Sea. Its continental position is disputed; while it is ge ...
. A serpulid worm is attached. File:OrhtocerasNautiloid092313.jpg, Silurian
Orthoceras ''Orthoceras'' ("straight horn") is a genus of extinct nautiloid cephalopod restricted to Middle Ordovician-aged marine limestones of the Baltic States and Sweden. This genus is sometimes called ''Orthoceratites''. Note it is sometimes misspelle ...
fossil File:Eocene fossil flower, Clare Family Florissant Fossil Quarry, Florissant, Colorado, USA - 20100807.jpg, Eocene fossil flower from Florissant, Colorado File:RoyLindmanMicraster.JPG, '' Micraster''
echinoid Sea urchins () are spiny, globular echinoderms in the class Echinoidea. About 950 species of sea urchin live on the seabed of every ocean and inhabit every depth zone from the intertidal seashore down to . The spherical, hard shells (tests) of ...
fossil from England File:Productid Permian Texas.JPG, Productid
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 ...
ventral valve; Roadian,
Guadalupian The Guadalupian is the second and middle series/epoch of the Permian. The Guadalupian was preceded by the Cisuralian and followed by the Lopingian. It is named after the Guadalupe Mountains of New Mexico and Texas, and dates between 272.95 ± ...
(Middle
Permian The Permian ( ) is a geologic period and System (stratigraphy), 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 ...
); Glass Mountains, Texas. File:Fossil agatized coral Florida.JPG, Agatized
coral Corals are marine invertebrates within the class Anthozoa of the phylum Cnidaria. They typically form compact colonies of many identical individual polyps. Coral species include the important reef builders that inhabit tropical oceans and ...
from the
Hawthorn Group The Hawthorn Group is a stratigraphic unit that includes several geologic formations of Late Oligocene to Pliocene age in North Florida, United States. It is known for its phosphate rock resources, and for its rich assemblages of Neogene vertebr ...
( Oligocene
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
),
Florida Florida is a state located in the Southeastern region of the United States. Florida is bordered to the west by the Gulf of Mexico, to the northwest by Alabama, to the north by Georgia, to the east by the Bahamas and Atlantic Ocean, and to ...
. An example of preservation by replacement. File:Fossils from Gotland beaches.jpg, Fossils from beaches of the
Baltic Sea The Baltic Sea is an arm of the Atlantic Ocean that is enclosed by Denmark, Estonia, Finland, Germany, Latvia, Lithuania, Poland, Russia, Sweden and the North and Central European Plain. The sea stretches from 53°N to 66°N latitude and ...
island of Gotland, placed on paper with 7 mm (0.28 inch) squares File:Dinosaur footprints in ToroToro Bolivia.jpg, Dinosaur footprints from Torotoro National Park in Bolivia.


See also

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References


Further reading


"Grand Canyon cliff collapse reveals 313 million-year-old fossil footprints"
21 August 2020, ''
CNN CNN (Cable News Network) is a multinational cable news channel headquartered in Atlanta, Georgia, U.S. Founded in 1980 by American media proprietor Ted Turner and Reese Schonfeld as a 24-hour cable news channel, and presently owned by ...
''
"Hints of fossil DNA discovered in dinosaur skull"
by Michael Greshko, 3 March 2020, '' National Geographic''
"Fossils for Kids , Learn all about how fossils are formed, the types of fossils and more!"
Video (2:23), 27 January 2020, ''Clarendon Learning''
"Fossil & their formation"
Video (9:55), 15 November 2019, ''
Khan Academy Khan Academy is an American non-profit educational organization created in 2008 by Sal Khan. Its goal is creating a set of online tools that help educate students. The organization produces short lessons in the form of videos. Its website also in ...
''
"How are dinosaur fossils formed?
by Lisa Hendry, ''
Natural History Museum, London The Natural History Museum in London is a museum that exhibits a vast range of specimens from various segments of natural history. It is one of three major museums on Exhibition Road in South Kensington, the others being the Science Museum an ...
''
"Fossils 101"
Video (4:27), 22 August 2019, '' National Geographic''
"How to Spot the Fossils Hiding in Plain Sight"
by Jessica Leigh Hester, 23 February 2018, ''
Atlas Obscura ''Atlas Obscura'' is an American-based online magazine and travel company. It was founded in 2009 by author Joshua Foer and documentary filmmaker/author Dylan Thuras. It catalogs unusual and obscure travel destinations via user-generated conten ...
''
"It's extremely hard to become a fossil"
, by
Olivia Judson Olivia P. Judson (born 1970) is a British evolutionary biologist and science writer. She is a former journalist for ''The Economist'', a former online columnist for ''The New York Times'' and has published in a number of other publications, includ ...
, 30 December 2008, ''
The New York Times ''The New York Times'' (''the Times'', ''NYT'', or the Gray Lady) is a daily newspaper based in New York City with a worldwide readership reported in 2020 to comprise a declining 840,000 paid print subscribers, and a growing 6 million paid d ...
''
"Bones Are Not the Only Fossils"
, by
Olivia Judson Olivia P. Judson (born 1970) is a British evolutionary biologist and science writer. She is a former journalist for ''The Economist'', a former online columnist for ''The New York Times'' and has published in a number of other publications, includ ...
, 4 March 2008, ''
The New York Times ''The New York Times'' (''the Times'', ''NYT'', or the Gray Lady) is a daily newspaper based in New York City with a worldwide readership reported in 2020 to comprise a declining 840,000 paid print subscribers, and a growing 6 million paid d ...
''


External links

*
The Virtual Fossil Museum throughout Time and Evolution

Paleoportal, geology and fossils of the United States

The Fossil Record, a complete listing of the families, orders, class and phyla found in the fossil record
* * * {{Authority control