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A fossil (from Classical Latin , ) 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 geochrono ...
. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, 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 Ma ...
, hair, petrified wood and DNA remnants. The totality of fossils is known as the ''fossil record''. Paleontology is the study of fossils: their age, method of formation, and evolutionary 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 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, 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 process ...
, and bioimmuration. Fossils vary in size from one- micrometre (1 µm) bacteria to dinosaurs 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 vertebrates, 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 adje ...
exoskeletons of invertebrates. Fossils may also consist of the marks left behind by the organism while it was alive, such as
animal tracks __notoc__ An animal track is an imprint left behind in soil, snow, or mud, or on some other ground surface, by an animal walking across it. Animal tracks are used by hunters in tracking their prey and by naturalists to identify animals living i ...
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 ...
s). These types of fossil are called trace fossils or ''ichnofossils'', as opposed to ''body fossils''. Some fossils are
biochemical Biochemistry or biological chemistry is the study of chemical processes within and relating to living organisms. A sub-discipline of both chemistry and biology, biochemistry may be divided into three fields: structural biology, enzymology and ...
and are called ''chemofossils'' or
biosignature A biosignature (sometimes called chemical fossil or molecular fossil) is any substance – such as an element, isotope, or molecule – or phenomenon that provides scientific evidence of past or present life. Measurable attribute ...
s.


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.


Fossilization processes

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


Permineralization

Permineralization 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. Minerals precipitate from the groundwater, occupying the empty spaces. This process can occur in very small spaces, such as within the
cell wall A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane. It can be tough, flexible, and sometimes rigid. It provides the cell with both structural support and protection, and also acts as a filtering me ...
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, 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 Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, biva ...
or snail or the hollow of a skull. 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, 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 concre ...
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 pr ...
to
calcite Calcite is a carbonate mineral and the most stable polymorph of calcium carbonate (CaCO3). It is a very common mineral, particularly as a component of limestone. Calcite defines hardness 3 on the Mohs scale of mineral hardness, based on scratc ...
.


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 sediment ...
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 (erythrocyt ...
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 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 Leeches are segmented parasitic or predatory worms that comprise the subclass Hirudinea within the phylum Annelida. They are closely related to the oligochaetes, which include the earthworm, and like them have soft, muscular segmented bodie ...
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.


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, which grows along a substrate, covering other sessile
sclerobiont Sclerobionts are collectively known as organisms living in or on any kind of hard substrate (Taylor and Wilson, 2003). A few examples of sclerobionts include ''Entobia'' borings, ''Gastrochaenolites'' borings, ''Talpina'' borings, serpulids, encrus ...
s. 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. The ...
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 sediments 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 Taxonomy (biology), 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 ...
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 fossils consist mainly of tracks and burrows, but also include
coprolite A coprolite (also known as a coprolith) is fossilized feces. Coprolites are classified as trace fossils as opposed to body fossils, as they give evidence for the animal's behaviour (in this case, diet) rather than morphology. The name is ...
s (fossil feces) 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. Th ...
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 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 The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ( ...
trace fossils including '' Rusophycus'', made by a trilobite File:Coprolite.jpg, A coprolite of a carnivorous dinosaur found in southwestern Saskatchewan 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 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 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) of larger animals or plants. Microfossils are of critical importance as a reservoir of
paleoclimate Paleoclimatology ( British spelling, palaeoclimatology) is the study of climates for which direct measurements were not taken. As instrumental records only span a tiny part of Earth's history, the reconstruction of ancient climate is important to ...
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 Ma ...
) is a natural polymer 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 which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
, though most dates to the
Cenozoic The Cenozoic ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterised by the dominance of mammals, birds and flowering plants, a cooling and drying climate, and the current configurat ...
. The excretion of the resin by certain plants is thought to be an evolutionary
adaptation In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the po ...
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 chordate ...
, 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 chitin, o ...
s such as insects and spiders, and only extremely rarely a vertebrate 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 p ...
. 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 ''Interna ...
. This will usually include "xylon" and a term indicating its presumed affinity, such as '' Araucarioxylon'' (wood of ''
Araucaria ''Araucaria'' (; original pronunciation: .ɾawˈka. ɾja is a genus of evergreen coniferous trees in the family Araucariaceae. There are 20 extant species in New Caledonia (where 14 species are endemic, see New Caledonian ''Araucaria ...
'' or some related genus), ''
Palmoxylon ''Palmoxylon'' (petrified palmwood) is an extinct genus of palm named from petrified wood found around the world. Fossil record This genus is known in the fossil record from the Late Cretaceous to the Miocene (from about 84.9 to 11.6 million y ...
'' (wood of an indeterminate palm), 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, or other biomolecules. Additionally, isotope 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. Subfossils are often found in depositionary environments, such as lake sediments, oceanic sediments, and soils. Once deposited, physical and chemical weathering can alter the state of preservation, and small subfossils can also be ingested by living organisms. Subfossil remains that date from the Mesozoic 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 species s ...
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, typica ...
carapace A carapace is a dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tortoises, the under ...
s, diatoms, and foraminifera. 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 age, and therefore are not old enough to be from the Pleistocene 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 the ...
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 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 sedimentary 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, 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 Bi ...
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 are commonly regarded as continents. Ordered from largest in area to smallest, these seven ...
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 mutations 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 nucleotid ...
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 The Cambrian explosion, Cambrian radiation, Cambrian diversification, or the Biological Big Bang refers to an interval of time approximately in the Cambrian Period when practically all major animal phyla started appearing in the fossil recor ...
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 Soft-bodied organisms are animals that lack skeletons. The group roughly corresponds to the group Vermes as proposed by Carl von Linné. All animals have muscles but, since muscles can only pull, never push, a number of animals have developed hard ...
with little to no role. It is replete with the mollusks, the vertebrates, the echinoderms, 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, wh ...
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 chitin, o ...
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 The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ( ...
period to the
present The present (or here'' and ''now) is the time that is associated with the events perceived directly and in the first time, not as a recollection (perceived more than once) or a speculation (predicted, hypothesis, uncertain). It is a period of ...
. Worldwide, some of the best examples of near-perfect fossilization are the
Cambrian The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ( ...
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 fos ...
, the Devonian
Hunsrück Slates The Hunsrück () is a long, triangular, pronounced upland in Rhineland-Palatinate, Germany. It is bounded by the valleys of the Moselle-Saar (north-to-west), the Nahe (south), and the Rhine (east). It is continued by the Taunus mountains, past ...
, the Jurassic Solnhofen limestone, and the
Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carboniferous ...
Mazon Creek 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 p ...
s of microorganisms, 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 Archea ...
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 Primordial may refer to: * Primordial era, an era after the Big Bang. See Chronology of the universe * Primordial sea (a.k.a. primordial ocean, ooze or soup). See Abiogenesis * Primordial nuclide, nuclides, a few radioactive, that formed before t ...
forms of the eukaryote chlorophytes (that is, green algae). One genus 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, san ...
is ''
Collenia ''Collenia'' is genus of fossil cyanobacteria that form a particular type of stromatolites. Description ''Collenia'' are stromatolites made up of convex layers flattened in the center, forming columnar colonies. The microorganisms involved wer ...
''. 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. The ...
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 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 cyanobacteria themselves reproduce asexually through cell division, they were instrumental in priming the environment for the evolutionary development of more complex eukaryotic 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 tem ...
Gammaproteobacteria) are thought to be largely responsible for increasing the amount of oxygen in the primeval earth's
atmosphere An atmosphere () is a layer of gas or layers of gases that envelop a planet, and is held in place by the gravity of the planetary body. A planet retains an atmosphere when the gravity is great and the temperature of the atmosphere is low. A s ...
through their continuing photosynthesis. Cyanobacteria use water, carbon dioxide and sunlight to create their food. A layer of mucus 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. 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 in ...
.
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 gr ...
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 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 Extraterrestrial life, colloquially referred to as alien life, is life that may occur outside Earth and which did not originate on Earth. No extraterrestrial life has yet been conclusively detected, although efforts are underway. Such life might ...
and thus could play an important role in the search for past or present life on the planet Mars. Furthermore, organic components (
biosignature A biosignature (sometimes called chemical fossil or molecular fossil) is any substance – such as an element, isotope, or molecule – or phenomenon that provides scientific evidence of past or present life. Measurable attribute ...
s) 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 be ...
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 Work ...
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 phototro ...
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 ...
microorganisms, as well as ancient water, including fluvio-lacustrine environments ( plains related to ancient rivers or lakes) that may have been habitable. 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 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 in ...
), and organic carbon on the planet Mars is now a primary NASA 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 n ...
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 agate Moss agate pebble, 1 inch (25 mm) long. Moss agate is a semi-precious gemstone formed from silicon dioxide. It is a form of chalcedony which includes minerals of a green color embedded in the stone, forming filaments and other patter ...
s, 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 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 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 foss ...
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 Taxonomy (biology), 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 ...
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 echinoderms 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 Neanderthals. 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 extant ...
, 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 in Roman mythology. Fossils appear to have directly contributed to the mythology of many civilizations, including the ancient Greeks. Classical Greek historian
Herodotos Herodotus ( ; grc, , }; BC) was an ancient Greek historian and geographer from the Greek city of Halicarnassus, part of the Persian Empire (now Bodrum, Turkey) and a later citizen of Thurii in modern Calabria (Italy). He is known for havin ...
wrote of an area near Hyperborea 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 back ...
s protected golden treasure. There was indeed gold mining in that approximate region, where beaked '' Protoceratops'' skulls were common as fossils. A later Greek scholar, Aristotle, 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 Persian polymath
Avicenna Ibn Sina ( fa, ابن سینا; 980 – June 1037 CE), commonly known in the West as Avicenna (), was a Persian polymath who is regarded as one of the most significant physicians, astronomers, philosophers, and writers of the Islamic ...
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 wrote of "
tongue stone Sharks continually shed their teeth; some Carcharhiniformes shed approximately 35,000 teeth in a lifetime, replacing those that fall out. There are four basic types of shark teeth: dense flattened, needle-like, pointed lower with triangular upp ...
s", which he called
glossopetra Sharks continually shed their teeth; some Carcharhiniformes shed approximately 35,000 teeth in a lifetime, replacing those that fall out. There are four basic types of shark teeth: dense flattened, needle-like, pointed lower with triangular upp ...
. 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
ammonite Ammonoids are a group of extinct marine mollusc animals in the subclass Ammonoidea of the class Cephalopoda. These molluscs, commonly referred to as ammonites, are more closely related to living coleoids (i.e., octopuses, squid and cuttlefish ...
s, 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 thic ...
'', a Cretaceous 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 ...
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 k ...
. 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, 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 cocain ...
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 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 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. 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 p ...
bamboo Bamboos are a diverse group of evergreen perennial flowering plants making up the subfamily Bambusoideae of the grass family Poaceae. Giant bamboos are the largest members of the grass family. The origin of the word "bamboo" is uncertain, but ...
s in the dry northern climate zone of what is now Yan'an, Shaanxi province, China, led him to advance early ideas of gradual climate change 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 intertwin ...
, 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 t ...
. 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 Empir ...
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 Ibn Sina ( fa, ابن سینا; 980 – June 1037 CE), commonly known in the West as Avicenna (), was a Persian polymath who is regarded as one of the most significant physicians, astronomers, philosophers, and writers of the Islamic ...
explained fossils' stoniness in '' The Book of Healing'': From the 13th century to the present day, scholars pointed out that the fossil skulls of
Deinotherium giganteum ''Deinotherium'' was a large elephant-like proboscidean that appeared in the Middle Miocene and survived until the Early Pleistocene. Although superficially resembling modern elephants, they had notably more flexible necks, limbs adapted to a mo ...
, 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, Cypru ...
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, 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 a ...
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, 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. Leonardo da Vinci 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 examined a shark, and made the association of its teeth with the "tongue stones" of ancient Greco-Roman mythology, concluding that those were not in fact the tongues of venomous snakes, but the teeth of some long-extinct species of shark. Robert Hooke (1635–1703) included micrographs of fossils in his '' Micrographia'' 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) 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 In geology, catastrophism theorises that the Earth has largely been shaped by sudden, short-lived, violent events, possibly worldwide in scope. This contrasts with uniformitarianism (sometimes called gradualism), according to which slow incremen ...
. 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'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 by ...
skeleton, sparked both public and scholarly interest.


Linnaeus and Darwin

Early naturalists well understood the similarities and differences of living species leading Linnaeus 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 The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ( ...
Period, 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 sessi ...
(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 Bi ...
or determining the age of rocks based on embedded fossils. For the first 150 years of geology, biostratigraphy and superposition were the only means for determining the relative age of rocks. The geologic time scale 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 directl ...
, 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 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, 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 am ...
or nucleotide 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'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 d ...
'' trilobite 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 ...
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  ...
tomographic analysis of early Cambrian bilaterian embryonic microfossils yielded new insights of metazoan 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 embryonic development. This nature has since been discovered to be extremely variable among Protostomia's me ...
, '' 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 chitin, o ...
development earlier than previously thought in the late Proterozoic. The preserved embryos from China and Siberia 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, Nematoda 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 chitin, o ...
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, 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 alum ...
activity or supply some essential minerals. The use of dinosaur bones as "dragon bones" has persisted in Traditional Chinese medicine 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 Dinosaurs are a divers ...
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 Ammonoids are a group of extinct marine mollusc animals in the subclass Ammonoidea of the class Cephalopoda. These molluscs, commonly referred to as ammonites, are more closely related to living coleoids (i.e., octopuses, squid and cuttlefish ...
fossils, each approximately 1.5 cm across File:Priscacara liops Green River Formation.jpg, Eocene 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 f ...
of Wyoming File:Trilobite2.jpg, A permineralized trilobite, ''
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 memb ...
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. File:The fossils from Cretaceous age found in Lebanon.jpg, Fossil shrimp ( Cretaceous) File:PetrifiedWood.jpg, Petrified wood in
Petrified Forest National Park Petrified Forest National Park is an American national park in Navajo and Apache counties in northeastern Arizona. Named for its large deposits of petrified wood, the park covers about , encompassing semi-desert shrub steppe as well as highly ero ...
, 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 conn ...
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 Cua ...
'' from Patagonia,
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 of Cyprus. A serpulid worm is attached. File:OrhtocerasNautiloid092313.jpg, Silurian Orthoceras 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, wh ...
ventral valve; Roadian, Guadalupian (Middle Permian); Glass Mountains, Texas. File:Fossil agatized coral Florida.JPG, Agatized coral from the Hawthorn Group ( OligoceneMiocene), Florida. 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 from ...
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 Torotoro National Park ( es, Parque Nacional Torotoro) is a national park and town in Bolivia. The town was founded about 250 years ago by the Quechuas and the Spanish. It is located in the eastern mountain ranges of the South American Andes co ...
in Bolivia.


See also

* * * * * * * * * *


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 t ...
''
"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''
"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, inclu ...
, 30 December 2008, '' The New York Times''
"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, inclu ...
, 4 March 2008, '' The New York Times''


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