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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 pe ...
, ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. 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, an ...
s, 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 M ...
, hair, petrified wood 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 fos ...
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 and the
relative ages Relative dating is the science of determining the relative order of past events (i.e., the age of an object in comparison to another), without necessarily determining their absolute age (i.e., estimated age). In geology, rock or superficial dep ...
of different fossils. The development of radiometric dating techniques in the early 20th century allowed scientists to quantitatively measure the
absolute ages 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 ...
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 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 ...
(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 taxon, taxa within the subphylum Vertebrata () (chordates with vertebral column, backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the ...
s, or the
chitin Chitin ( C8 H13 O5 N)n ( ) is a long-chain polymer of ''N''-acetylglucosamine, an amide derivative of glucose. Chitin is probably the second most abundant polysaccharide in nature (behind only cellulose); an estimated 1 billion tons of chit ...
ous 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 a ...
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 __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 livi ...
or
feces Feces ( or faeces), known colloquially and in slang as poo and poop, are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the large intestine. Feces contain a rela ...
( coprolites). These types of fossil are called trace fossils 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 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 tota ...
.


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 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 unconsolidat ...
. 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 mec ...
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 cuticle, animal coverings, such as the arthropod exoskeleton, have diffe ...
, feathers or even soft tissues. This is a form of diagenesis.


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, 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, biv ...
or snail 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, t ...
. 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 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 to calcite.


Adpression (compression-impression)

Compression fossils, 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. Often what remains is a carbonaceous film 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-aFeO(OH)) associated with soft tissues recovered from dinosaur fossils. Based on various experiments that studied the interaction of iron in haemoglobin with blood vessel tissue they proposed that solution hypoxia coupled with iron chelation 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 Geological period, geologic period and System (stratigraphy), 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 J ...
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 b ...
from the Silurian Waukesha Biota of Wisconsin. File:Lycopod axis.jpg, Partially coalified axis (branch) of a lycopod 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 skeletal organism, such as a bryozoan or an oyster, which grows along a
substrate Substrate may refer to: Physical layers *Substrate (biology), the natural environment in which an organism lives, or the surface or medium on which an organism grows or is attached ** Substrate (locomotion), the surface over which an organism lo ...
, 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. ...
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 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 ...
(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 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 coprolites (fossil
feces Feces ( or faeces), known colloquially and in slang as poo and poop, are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the large intestine. Feces contain a rela ...
) and marks left by feeding. Trace fossils are particularly significant because they represent a data source that is not limited to animals with easily 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. ...
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 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 ag ...
trace fossils including '' Rusophycus'', made by a trilobite 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 ...
File:Climactichnites wilsoni, densely packed.jpg, Densely packed, subaerial or nearshore trackways ('' Climactichnites 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 es ...
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 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 coccolithophores) 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 t ...
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 M ...
) 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 and ...
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 ...
. 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 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 chordate ...
, predominantly
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
s such as insects and spiders, and only extremely rarely a
vertebrate Vertebrates () comprise all animal taxon, taxa within the subphylum Vertebrata () (chordates with vertebral column, backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the ...
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. 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 '' Inte ...
. This will usually include "xylon" and a term indicating its presumed affinity, such as '' Araucarioxylon'' (wood of '' Araucaria'' or some related genus), '' Palmoxylon'' (wood of an indeterminate palm), or ''Castanoxylon'' (wood of an indeterminate
chinkapin Chinquapin or chinkapin may refer to: Plants * Chinquapin or chinkapin, any of the shrubs in the genus '' Castanopsis'' * Chinquapin or chinkapin, any of the several trees and shrubs in the genus ''Chrysolepis'' * Chinquapin or chinkapin, some of ...
).


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 Radiocarbon dating (also referred to as carbon dating or carbon-14 dating) is a method for determining the age of an object containing organic material by using the properties of radiocarbon, a radioactive isotope of carbon. The method was de ...
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, respon ...
, 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 number ...
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 movemen ...
can alter the state of preservation, and small subfossils can also be ingested by living
organisms In biology, an organism () is any living system that functions as an individual entity. All organisms are composed of cells ( cell theory). Organisms are classified by taxonomy into groups such as multicellular animals, plants, and f ...
. 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 Creta ...
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 head capsules, ostracod carapaces, 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 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 ...
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 zircons 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. 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 Stratigraphy is a branch of geology concerned with the study of rock layers ( strata) and layering (stratification). It is primarily used in the study of sedimentary and layered volcanic rocks. Stratigraphy has three related subfields: lithost ...
to date fossils. Stratigraphy is the science of deciphering the "layer-cake" that is the
sediment Sediment is a naturally occurring material that is broken down by processes of weathering and erosion, and is subsequently transported by the action of wind, water, or ice or by the force of gravity acting on the particles. For example, 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 di ...
, 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 fossils 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
mutation In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, m ...
s accumulate at a constant rate. These "
molecular clock The molecular clock is a figurative term for a technique that uses the mutation rate of biomolecules to deduce the time in prehistory when two or more life forms diverged. The biomolecular data used for such calculations are usually nucleo ...
s", however, are fallible, and provide only 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 taxon, taxa within the subphylum Vertebrata () (chordates with vertebral column, backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the ...
s, 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, ...
s and some groups of
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
s.


Sites


Lagerstätten

Fossil sites with exceptional preservation—sometimes including preserved soft tissues—are known as Lagerstätten—German for "storage places". These formations may have resulted from carcass burial in an anoxic 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 ag ...
period to the present. 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 ag ...
Maotianshan shales and Burgess Shale, 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 The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), 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 J ...
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 concre ...
localities.


Stromatolites

Stromatolites are layered accretionary structures formed in shallow water by the trapping, binding and cementation of sedimentary grains by biofilms 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 eon (geology), geologic eons of History of Earth, Earth's history, representing the time from . The Archean was preceded by the Hadean Eon and followed by the Proterozo ...
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 The Proterozoic () is a geological eon spanning the time interval from 2500 to 538.8million years ago. It is the most recent part of the Precambrian "supereon". It is also the longest eon of the Earth's geologic time scale, and it is subdivided ...
) 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 ...
forms of the
eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bact ...
chlorophytes Chlorophyta or Prasinophyta is a taxon of green algae informally called chlorophytes. The name is used in two very different senses, so care is needed to determine the use by a particular author. In older classification systems, it refers to ...
(that is,
green algae The green algae (singular: green alga) are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta and Charophyta/ Streptophyta. The land plants ( Embryophytes) have emerged deep in the Charophyte alg ...
). 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 n ...
of stromatolite very common in the geologic record 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), 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. ...
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 rap ...
; 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 tempe ...
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 we ...
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. ...
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, transparent, tasteless, odorless, and nearly colorless chemical substance, which is the main constituent of Earth's hydrosphere and the fluids of all known living organisms (in which it acts as ...
,
carbon dioxide Carbon dioxide ( chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is t ...
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 wh ...
. 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 oncolites 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 ...
. Thrombolites 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 e ...
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. Furthermore, organic components ( biosignatures) that are often associated with biominerals are believed to play crucial roles in both pre-biotic and
biotic Biotics describe living or once living components of a community; for example organisms, such as animals and plants. Biotic may refer to: *Life, the condition of living organisms *Biology, the study of life * Biotic material, which is derived from ...
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 Works", ...
ic,
chemotroph A Chemotroph is an organism that obtains energy by the oxidation of electron donors in their environments. These molecules can be organic molecule, organic (chemoorganotrophs) or inorganic compound, inorganic (chemolithotrophs). The chemotroph de ...
ic and/or chemolithoautotrophic
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 Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salts and other total dissolved solids. Although the ...
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 lar ...
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, taphonomy (related to fossils), and organic carbon on the planet Mars is now a primary
NASA The National Aeronautics and Space Administration (NASA ) is an independent agency of the US federal government responsible for the civil space program, aeronautics research, and space research. NASA was established in 1958, succeedi ...
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 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 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 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 The Paleolithic or Palaeolithic (), also called the Old Stone Age (from Greek: παλαιός '' palaios'', "old" and λίθος ''lithos'', "stone"), is a period in human prehistory that is distinguished by the original development of stone to ...
stone knives in Europe, with fossil echinoderms set precisely at the hand grip, going all the way back to '' Homo heidelbergensis'' 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 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 exta ...
, 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, 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 where gryphons protected golden treasure. There was indeed gold mining in that approximate region, where beaked '' Protoceratops'' 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 Greece, Classical period in Ancient Greece. Taught by Plato, he was the founder of the Peripatet ...
, 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. H ...
ous exhalations, 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 Turkme ...
n 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 Islam ...
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, 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 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 n ...
, 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 toadstones, 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'', 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 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. 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 aphrodisiacs. 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 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 Shen Kuo hypothesized that marine fossils found in a stratum, geological stratum of mountains located hundreds of miles from the Pacific Ocean was evidence that a prehistoric seashore had once existed there and Geomorphology, 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 bamboos 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 (general concept), 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, fossilized sea creatures on mountainsides were seen as proof of the biblical deluge of Noah's Ark. After observing the existence of seashells in mountains, the List of ancient Greek philosophers, ancient Greek philosopher Xenophanes (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 people, Persian
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 Islam ...
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, found in Crete and Greece, might have been interpreted as being the skulls of the Cyclopes 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 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 Thunderstone (folklore)#Fossils as thunderstones, 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 loaf, 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 forams. 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 (geologist), 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. 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 skeleton, sparked both public and scholarly interest.


Linnaeus and Darwin

Early Natural science, naturalists well understood the similarities and differences of living species leading Carl Linnaeus, 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, 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 ag ...
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. phylum, 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 Micropaleontology, microfossils. However, macroscopic fossils are now known from the late Proterozoic. The Ediacara biota (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 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 dating, 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 Stratigraphy, stratigraphers. Since the early years of the twentieth century, absolute dating methods, such as radiometric dating (including K–Ar dating, potassium/argon, Argon–argon dating, argon/argon, uranium–lead dating, uranium series, and, for very recent fossils,
radiocarbon dating Radiocarbon dating (also referred to as carbon dating or carbon-14 dating) is a method for determining the age of an object containing organic material by using the properties of radiocarbon, a radioactive isotope of carbon. The method was de ...
) 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 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'' 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 1971. Synchrotron X-ray Tomography, tomographic analysis of early Cambrian bilaterian embryonic microfossils yielded new insights of Animal, 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'' and a putative, primitive protostome, ''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 Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
development earlier than previously thought in the late Proterozoic. The preserved embryos from China and Siberia underwent rapid Diagenesis, diagenetic 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, Nematode, Nematoda and Arthropoda. 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 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 and include the use of fossils as talismans. 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 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 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, HP, India File:Amonite Cropped.jpg, Three small ammonite fossils, each approximately 1.5 cm across File:Priscacara liops Green River Formation.jpg, Eocene fossil fish ''Priscacara liops'' from the Green River Formation of Wyoming File:Trilobite2.jpg, A permineralized trilobite, ''Asaphus kowalewskii'' File:Carcharodontosaurus and Megalodon teeth.jpg, Megalodon and ''Carcharodontosaurus'' teeth. The latter was found in the Sahara Desert. 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 Conifer cone, cone of ''Araucaria mirabilis'' from Patagonia, Argentina dating from the Jurassic Period (approx. 210 Ma (unit), Ma) File:CyprusPlioceneGastropod.JPG, A fossil gastropod from the Pliocene of Cyprus. A Serpulidae, 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 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, ...
ventral valve; Roadian, Guadalupian (Middle Permian); Glass Mountains, Texas. File:Fossil agatized coral Florida.JPG, Agate, Agatized coral from the Hawthorn Group (Oligocene–Miocene), Florida. An example of preservation by replacement. File:Fossils from Gotland beaches.jpg, Fossils from beaches of the Baltic Sea 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''
"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''
"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''
"It's extremely hard to become a fossil"
, by Olivia Judson, 30 December 2008, ''The New York Times''
"Bones Are Not the Only Fossils"
, by Olivia Judson, 4 March 2008, ''The New York Times''


External links

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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 Fossils,