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A fossil (from
Classical Latin Classical Latin is the form of Literary Latin recognized as a literary standard by writers of the late Roman Republic and early Roman Empire. It was used from 75 BC to the 3rd century AD, when it developed into Late Latin. In some later period ...
, ) is any preserved remains, impression, or trace of any once-living thing from a past
geological age The geologic time scale, or geological time scale, (GTS) is a representation of time based on the rock record of Earth. It is a system of chronological dating that uses chronostratigraphy (the process of relating strata to time) and geochrono ...
. Examples include
bone A bone is a rigid organ that constitutes part of the skeleton in most vertebrate animals. Bones protect the various other organs of the body, produce red and white blood cells, store minerals, provide structure and support for the body, ...
s, shells, exoskeletons, 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 Hair is a protein filament that grows from follicles found in the dermis. Hair is one of the defining characteristics of mammals. The human body, apart from areas of glabrous skin, is covered in follicles which produce thick terminal and fi ...
, 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 fossi ...
is the study of fossils: their age, method of formation, and
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
ary significance. Specimens are usually considered to be fossils if they are over 10,000 years old. The oldest fossils are around 3.48 billion years old to 4.1 billion years old. Early edition, published online before print. The observation in the 19th century that certain fossils were associated with certain rock
strata In geology and related fields, a stratum ( : strata) is a layer of rock or sediment characterized by certain lithologic properties or attributes that distinguish it from adjacent layers from which it is separated by visible surfaces known as e ...
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 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 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 accurac ...
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, and bioimmuration. Fossils vary in size from one-
micrometre The micrometre (American and British English spelling differences#-re, -er, international spelling as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer (American and British English spelling differences# ...
(1 µm) bacteria to
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
s and trees, many meters long and weighing many tons. A fossil normally preserves only a portion of the deceased organism, usually that portion that was partially mineralized during life, such as the bones and teeth of
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with ...
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 ad ...
exoskeletons of
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chorda ...
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 livin ...
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 ...
(
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 an ...
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 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 to ...
.


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 Pore space in soil, soil pore spaces and in the fractures of stratum, rock formations. About 30 percent of all readily available freshwater in the world is groundwater. A unit ...
. 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 Plant cells are the cells present in green plants, photosynthetic eukaryotes of the kingdom Plantae. Their distinctive features include primary cell walls containing cellulose, hemicelluloses and pectin, the presence of plastids with the capab ...
. Small scale permineralization can produce very detailed fossils. For permineralization to occur, the organism must become covered by sediment soon after death, otherwise the remains are destroyed by scavengers or decomposition. The degree to which the remains are decayed when covered determines the later details of the fossil. Some fossils consist only of skeletal remains or teeth; other fossils contain traces of
skin Skin is the layer of usually soft, flexible outer tissue covering the body of a vertebrate animal, with three main functions: protection, regulation, and sensation. Other animal coverings, such as the arthropod exoskeleton, have different ...
,
feather Feathers are epidermal growths that form a distinctive outer covering, or plumage, on both avian (bird) and some non-avian dinosaurs and other archosaurs. They are the most complex integumentary structures found in vertebrates and a premie ...
s 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 (biology), class of marine and freshwater Mollusca, molluscs that have laterally compressed bodies enclosed by a shell consisting of two hing ...
or
snail A snail is, in loose terms, a shelled gastropod. The name is most often applied to land snails, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class ...
or the hollow of a
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
. Endocasts are sometimes termed , especially when bivalves are preserved this way.


Authigenic mineralization

This is a special form of cast and mold formation. If the chemistry is right, the organism (or fragment of organism) can act as a nucleus for the precipitation of minerals such as siderite, 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 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 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 The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, wh ...
to the
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
found that reasonably well-preserved fibrils that probably represent
collagen Collagen () is the main structural protein in the extracellular matrix found in the body's various connective tissues. As the main component of connective tissue, it is the most abundant protein in mammals, making up from 25% to 35% of the whol ...
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 parasitism, parasitic or Predation, predatory worms that comprise the Class (biology), subclass Hirudinea within the phylum Annelida. They are closely related to the Oligochaeta, oligochaetes, which include the earthwor ...
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 Oyster is the common name for a number of different families of salt-water bivalve molluscs that live in marine or brackish habitats. In some species, the valves are highly calcified, and many are somewhat irregular in shape. Many, but not ...
, which grows along a substrate, covering other sessile
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. T ...
to the Recent.


Types


Index

Index fossils (also known as guide fossils, indicator fossils or zone fossils) are fossils used to define and identify geologic periods (or faunal stages). They work on the premise that, although different
sediment Sediment is a naturally occurring material that is broken down by processes of weathering and erosion, and is subsequently transported by the action of wind, water, or ice or by the force of gravity acting on the particles. For example, sand ...
s may look different depending on the conditions under which they were deposited, they may include the remains of the same
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
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 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. T ...
s). Coprolites are classified as trace fossils as opposed to body fossils, as they give evidence for the animal's behaviour (in this case, diet) rather than morphology. They were first described by William Buckland 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 ''Rusophycus'' is a trace fossil ichnogenus allied to '' Cruziana''. ''Rusophycus'' is the resting trace, recording the outline of the tracemaker; ''Cruziana'' is made when the organism moved. The sculpture of ''Rusophycus'' may reveal the appr ...
'', made by a
trilobite Trilobites (; meaning "three lobes") are extinct marine arthropods that form the class Trilobita. Trilobites form one of the earliest-known groups of arthropods. The first appearance of trilobites in the fossil record defines the base of the ...
File:Coprolite.jpg, A coprolite of a carnivorous dinosaur found in southwestern
Saskatchewan Saskatchewan ( ; ) is a Provinces and territories of Canada, province in Western Canada, 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 t ...
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 e ...
on a Cambrian tidal flat


Transitional

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


Microfossils

Microfossil is a descriptive term applied to fossilized plants and animals whose size is just at or below the level at which the fossil can be analyzed by the naked eye. A commonly applied cutoff point between "micro" and "macro" fossils is 1 mm. Microfossils may either be complete (or near-complete) organisms in themselves (such as the marine plankters
foraminifera Foraminifera (; Latin for "hole bearers"; informally called "forams") are single-celled organisms, members of a phylum or class of amoeboid protists characterized by streaming granular ectoplasm for catching food and other uses; and commonly ...
and
coccolithophore Coccolithophores, or coccolithophorids, are single celled organisms which are part of the phytoplankton, the autotrophic (self-feeding) component of the plankton community. They form a group of about 200 species, and belong either to the king ...
s) or component parts (such as small teeth or spores) 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 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 a ...
found in many types of strata throughout the world, even the
Arctic The Arctic ( or ) is a polar regions of Earth, 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 (Greenla ...
. 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 per ...
, 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 configu ...
. 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 chordat ...
, predominantly
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
s such as insects and spiders, and only extremely rarely a
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with ...
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'' (wood of an indeterminate palm), or ''Castanoxylon'' (wood of an indeterminate chinkapin).


Subfossil

The term
subfossil 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 ...
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, res ...
, or other biomolecules. Additionally,
isotope Isotopes are two or more types of atoms that have the same atomic number (number of protons in their nuclei) and position in the periodic table (and hence belong to the same chemical element), and that differ in nucleon numbers ( mass num ...
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 (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 ...
. Subfossils are often found in depositionary environments, such as lake sediments, oceanic sediments, and soils. Once deposited, physical and chemical
weathering Weathering is the deterioration of rocks, soils and minerals as well as wood and artificial materials through contact with water, atmospheric gases, and biological organisms. Weathering occurs '' in situ'' (on site, with little or no movement ...
can alter the state of preservation, and small subfossils can also be ingested by living
organisms In biology, an organism () is any living system that functions as an individual entity. All organisms are composed of cells ( cell theory). Organisms are classified by taxonomy into groups such as multicellular animals, plants, and fu ...
. Subfossil remains that date from the
Mesozoic The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretace ...
are exceptionally rare, are usually in an advanced state of decay, and are consequently much disputed. The vast bulk of subfossil material comes from
Quaternary The Quaternary ( ) is the current and most recent of the three periods of the Cenozoic Era in the geologic time scale of the International Commission on Stratigraphy (ICS). It follows the Neogene Period and spans from 2.58 million year ...
sediments, including many subfossilized
chironomid The Chironomidae (informally known as chironomids, nonbiting midges, or lake flies) comprise a family of nematoceran flies with a global distribution. They are closely related to the Ceratopogonidae, Simuliidae, and Thaumaleidae. Many specie ...
head capsules,
ostracod Ostracods, or ostracodes, are a class of the Crustacea (class Ostracoda), sometimes known as seed shrimp. Some 70,000 species (only 13,000 of which are extant) have been identified, grouped into several orders. They are small crustaceans, typic ...
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 unde ...
s,
diatoms A diatom (Neo-Latin ''diatoma''), "a cutting through, a severance", from el, διάτομος, diátomos, "cut in half, divided equally" from el, διατέμνω, diatémno, "to cut in twain". is any member of a large group comprising sev ...
, and
foraminifera Foraminifera (; Latin for "hole bearers"; informally called "forams") are single-celled organisms, members of a phylum or class of amoeboid protists characterized by streaming granular ectoplasm for catching food and other uses; and commonly ...
. For remains such as molluscan
seashell A seashell or sea shell, also known simply as a shell, is a hard, protective outer layer usually created by an animal or organism that lives in the sea. The shell is part of the body of the animal. Empty seashells are often found washe ...
s, 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 (geology), 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 fina ...
epoch.


Chemical fossils

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


Dating


Estimating dates

Paleontology seeks to map out how life evolved across geologic time. A substantial hurdle is the difficulty of working out fossil ages. Beds that preserve fossils typically lack the radioactive elements needed for
radiometric dating Radiometric dating, radioactive dating or radioisotope dating is a technique which is used to date materials such as rocks or carbon, in which trace radioactive impurities were selectively incorporated when they were formed. The method compares ...
. This technique is our only means of giving rocks greater than about 50 million years old an absolute age, and can be accurate to within 0.5% or better. Although radiometric dating requires careful laboratory work, its basic principle is simple: the rates at which various radioactive elements decay 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, sand ...
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 d ...
, 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 The Cambrian explosion, Cambrian radiation, Cambrian diversification, or the Biological Big Bang refers to an interval of time approximately in the Cambrian Period when practically all major animal phyla started appearing in the fossil record. ...
first evolved, and estimates produced by different techniques may vary by a factor of two.


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 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 esti ...
, the
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with ...
s, the
echinoderm An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the ...
s, the brachiopods and some groups of
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
s.


Sites


Lagerstätten

Fossil sites with exceptional preservation—sometimes including preserved soft tissues—are known as
Lagerstätte A Lagerstätte (, from '' Lager'' 'storage, lair' '' Stätte'' 'place'; plural ''Lagerstätten'') is a sedimentary deposit that exhibits extraordinary fossils with exceptional preservation—sometimes including preserved soft tissues. These f ...
n—German for "storage places". These formations may have resulted from carcass burial in an
anoxic The term anoxia means a total depletion in the level of oxygen, an extreme form of hypoxia or "low oxygen". The terms anoxia and hypoxia are used in various contexts: * Anoxic waters, sea water, fresh water or groundwater that are depleted of diss ...
environment with minimal bacteria, thus slowing decomposition. Lagerstätten span geological time from the
Cambrian 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 and Burgess Shale, the
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, wh ...
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 geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
Solnhofen limestone, and the
Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carboniferou ...
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 concretio ...
localities.


Stromatolites

Stromatolites are layered accretionary
structure A structure is an arrangement and organization of interrelated elements in a material object or system, or the object or system so organized. Material structures include man-made objects such as buildings and machines and natural objects such a ...
s 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 po ...
s of
microorganism A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in old ...
s, especially
cyanobacteria Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, bl ...
. Stromatolites provide some of the most ancient fossil records of life on Earth, dating back more than 3.5 billion years ago. Stromatolites were much more abundant in Precambrian times. While older,
Archean The Archean Eon ( , also spelled Archaean or Archæan) is the second of four geologic eons of Earth's history, representing the time from . The Archean was preceded by the Hadean Eon and followed by the Proterozoic. The Earth during the Arc ...
fossil remains are presumed to be
colonies In modern parlance, a colony is a territory subject to a form of foreign rule. Though dominated by the foreign colonizers, colonies remain separate from the administration of the original country of the colonizers, the '' metropolitan state'' ...
of
cyanobacteria Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, bl ...
, 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 i ...
) 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 Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacter ...
chlorophytes (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 alga ...
). One
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of stromatolite very common in the geologic record 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. T ...
evolutionary radiation; 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 Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
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 Gammaproteobacteria is a class of bacteria in the phylum Pseudomonadota (synonym Proteobacteria). It contains about 250 genera, which makes it the most genera-rich taxon of the Prokaryotes. Several medically, ecologically, and scientifically imp ...
) 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 ...
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 ...
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 in ...
. 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 Sunlight is a portion of the electromagnetic radiation given off by the Sun, in particular infrared, visible, and ultraviolet light. On Earth, sunlight is scattered and filtered through Earth's atmosphere, and is obvious as daylight when ...
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 w ...
. 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. 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 ea ...
provides an extremely well exposed example of the thrombolite-stromatolite-metazoan reefs that developed during the Proterozoic period, the stromatolites here being better developed in updip locations under conditions of higher current velocities and greater sediment influx.


Astrobiology

It has been suggested that biominerals could be important indicators of
extraterrestrial life 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 Mars is the fourth planet from the Sun and the second-smallest planet in the Solar System, only being larger than Mercury. In the English language, Mars is named for the Roman god of war. Mars is a terrestrial planet with a thin at ...
. Furthermore, organic components (
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 ...
based on
autotroph An autotroph or primary producer is an organism that produces complex organic compounds (such as carbohydrates, fats, and proteins) using carbon from simple substances such as carbon dioxide,Morris, J. et al. (2019). "Biology: How Life Wo ...
ic, chemotrophic and/or
chemolithoautotrophic A lithoautotroph is an organism which derives energy from reactions of reduced compounds of mineral (inorganic) origin. Two types of lithoautotrophs are distinguished by their energy source; photolithoautotrophs derive their energy from light while ...
microorganism A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in old ...
s, as well as ancient water, including fluvio-lacustrine environments (
plain In geography, a plain is a flat expanse of land that generally does not change much in elevation, and is primarily treeless. Plains occur as lowlands along valleys or at the base of mountains, as coastal plains, and as plateaus or uplands ...
s related to ancient
river A river is a natural flowing watercourse, usually freshwater, flowing towards an ocean, sea, lake or another river. In some cases, a river flows into the ground and becomes dry at the end of its course without reaching another body of ...
s or
lake A lake is an area filled with water, localized in a basin, surrounded by land, and distinct from any river or other outlet that serves to feed or drain the lake. Lakes lie on land and are not part of the ocean, although, like the much large ...
s) that may have been habitable. The search for evidence of habitability, taphonomy (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 ...
), and organic carbon on the planet
Mars Mars is the fourth planet from the Sun and the second-smallest planet in the Solar System, only being larger than Mercury. In the English language, Mars is named for the Roman god of war. Mars is a terrestrial planet with a thin at ...
is now a primary
NASA The National Aeronautics and Space Administration (NASA ) is an independent agency of the US federal government responsible for the civil space program, aeronautics research, and space research. NASA was established in 1958, succeedin ...
objective.


Pseudofossils

''Pseudofossils'' are visual patterns in rocks that are produced by geologic processes rather than biologic processes. They can easily be mistaken for real fossils. Some pseudofossils, such as geological
dendrite Dendrites (from Greek δένδρον ''déndron'', "tree"), also dendrons, are branched protoplasmic extensions of a nerve cell that propagate the electrochemical stimulation received from other neural cells to the cell body, or soma, of the ...
crystals, are formed by naturally occurring fissures in the rock that get filled up by percolating minerals. Other types of pseudofossils are kidney ore (round shapes in iron ore) and
moss 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 fossi ...
s examine the fossil record to understand the process of evolution and the way particular
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
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 too ...
stone knives in Europe, with fossil
echinoderm An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the ...
s set precisely at the hand grip, going all the way back to '' Homo heidelbergensis'' and
Neanderthal Neanderthals (, also ''Homo neanderthalensis'' and erroneously ''Homo sapiens neanderthalensis''), also written as Neandertals, are an Extinction, extinct species or subspecies of archaic humans who lived in Eurasia until about 40,000 years ag ...
s. These ancient peoples also drilled holes through the center of those round fossil shells, apparently using them as beads for necklaces. The ancient Egyptians gathered fossils of species that resembled the bones of modern species they worshipped. The god
Set Set, The Set, SET or SETS may refer to: Science, technology, and mathematics Mathematics *Set (mathematics), a collection of elements *Category of sets, the category whose objects and morphisms are sets and total functions, respectively Electro ...
was associated with the
hippopotamus The hippopotamus ( ; : hippopotamuses or hippopotami; ''Hippopotamus amphibius''), also called the hippo, common hippopotamus, or river hippopotamus, is a large semiaquatic mammal native to sub-Saharan Africa. It is one of only two 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 scholar,
Aristotle Aristotle (; grc-gre, Ἀριστοτέλης ''Aristotélēs'', ; 384–322 BC) was a Greek philosopher and polymath during the Classical period in Ancient Greece. Taught by Plato, he was the founder of the Peripatetic school of ...
, 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 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 Turkmeni ...
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 Islamic ...
modified into the theory of petrifying
fluid In physics, a fluid is a liquid, gas, or other material that continuously deforms (''flows'') under an applied shear stress, or external force. They have zero shear modulus, or, in simpler terms, are substances which cannot resist any shear ...
s (). Recognition of fossil seashells as originating in the sea was built upon in the 14th century by
Albert of Saxony en, Frederick Augustus Albert Anthony Ferdinand Joseph Charles Maria Baptist Nepomuk William Xavier George Fidelis , image = Albert of Saxony by Nicola Perscheid c1900.jpg , image_size = , caption = Photograph by Nicola Persch ...
, and accepted in some form by most naturalists by the 16th century. Roman naturalist
Pliny the Elder Gaius Plinius Secundus (AD 23/2479), called Pliny the Elder (), was a Roman author, naturalist and natural philosopher, and naval and army commander of the early Roman Empire, and a friend of the emperor Vespasian. He wrote the encyclopedic ' ...
wrote of " tongue stones", which he called glossopetra. These were fossil shark teeth, thought by some classical cultures to look like the tongues of people or snakes. He also wrote about the
horns of Ammon The horns of Ammon were curling ram horns, used as a symbol of the Egyptian deity Ammon (also spelled Amun or Amon). Because of the visual similarity, they were also associated with the fossils shells of ancient snails and cephalopods, the latter ...
, which are fossil
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 cuttlefis ...
s, 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 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. 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 ''Homo erectus'' (; meaning "upright man") is an extinct species of archaic human from the Pleistocene, with its earliest occurrence about 2 million years ago. Several human species, such as '' H. heidelbergensis'' and '' H. antecessor ...
'' were often mistaken for "
dragon A dragon is a reptilian legendary creature that appears in the folklore of many cultures worldwide. Beliefs about dragons vary considerably through regions, but dragons in western cultures since the High Middle Ages have often been depicted a ...
bones" and used as medicine and
aphrodisiac An aphrodisiac is a substance that increases sexual desire, sexual attraction, sexual pleasure, or sexual behavior. Substances range from a variety of plants, spices, foods, and synthetic chemicals. Natural aphrodisiacs like cannabis or cocai ...
s. In addition, some of these fossil bones are collected as "art" by scholars, who left scripts on various artifacts, indicating the time they were added to a collection. One good example is the famous scholar Huang Tingjian of the
Song Dynasty The Song dynasty (; ; 960–1279) was an imperial dynasty of China that began in 960 and lasted until 1279. The dynasty was founded by Emperor Taizu of Song following his usurpation of the throne of the Later Zhou. The Song conquered the res ...
during the 11th century, who kept a specific seashell fossil with his own poem engraved on it. In his ''
Dream Pool Essays ''The Dream Pool Essays'' (or ''Dream Torrent Essays'') was an extensive book written by the Chinese polymath and statesman Shen Kuo (1031–1095), published in 1088 during the Song dynasty (960–1279) of China. Shen compiled this encycloped ...
'' 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 Shen Kuo (; 1031–1095) or Shen Gua, courtesy name Cunzhong (存中) and pseudonym Mengqi (now usually given as Mengxi) Weng (夢溪翁),Yao (2003), 544. was a Chinese polymathic scientist and statesman of the Song dynasty (960–1279). Shen wa ...
hypothesized that marine fossils found in a geological stratum of mountains located hundreds of miles from the
Pacific Ocean The Pacific Ocean is the largest and deepest of Earth's five oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean (or, depending on definition, to Antarctica) in the south, and is bounded by the conti ...
was evidence that a prehistoric seashore had once existed there and shifted over centuries of time.Needham, Joseph. (1959). ''Science and Civilization in China: Volume 3, Mathematics and the Sciences of the Heavens and the Earth''.
Cambridge University Press Cambridge University Press is the university press of the University of Cambridge. Granted letters patent by King Henry VIII in 1534, it is the oldest university press in the world. It is also the King's Printer. Cambridge University Pr ...
. 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, ...
s in the dry northern climate zone of what is now
Yan'an Yan'an (; ), alternatively spelled as Yenan is a prefecture-level city in the Shaanbei region of Shaanxi province, China, bordering Shanxi to the east and Gansu to the west. It administers several counties, including Zhidan (formerly Bao'an) ...
,
Shaanxi Shaanxi (alternatively Shensi, see § Name) is a landlocked province of China. Officially part of Northwest China, it borders the province-level divisions of Shanxi (NE, E), Henan (E), Hubei (SE), Chongqing (S), Sichuan (SW), Gansu (W), N ...
province, China, led him to advance early ideas of gradual
climate change In common usage, climate change describes global warming—the ongoing increase in global average temperature—and its effects on Earth's climate system. Climate change in a broader sense also includes previous long-term changes to ...
due to bamboo naturally growing in wetter climate areas.Rafferty, John P. (2012). ''Geological Sciences; Geology: Landforms, Minerals, and Rocks''. New York: Britannica Educational Publishing, p. 6. In medieval
Christendom Christendom historically refers to the Christian states, Christian-majority countries and the countries in which Christianity dominates, prevails,SeMerriam-Webster.com : dictionary, "Christendom"/ref> or is culturally or historically 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 ...
. After observing the existence of seashells in mountains, the ancient Greek philosopher
Xenophanes Xenophanes of Colophon (; grc, Ξενοφάνης ὁ Κολοφώνιος ; c. 570 – c. 478 BC) was a Greek philosopher, theologian, poet, and critic of Homer from Ionia who travelled throughout the Greek-speaking world in early Classica ...
(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
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, found in
Crete Crete ( el, Κρήτη, translit=, Modern: , Ancient: ) is the largest and most populous of the Greek islands, the 88th largest island in the world and the fifth largest island in the Mediterranean Sea, after Sicily, Sardinia, Cyprus, ...
and Greece, might have been interpreted as being the skulls of the Cyclopes of
Greek mythology A major branch of classical mythology, Greek mythology is the body of myths originally told by the ancient Greeks, and a genre of Ancient Greek folklore. These stories concern the origin and nature of the world, the lives and activities o ...
, 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 Elephantida ...
cousins, though in fact it's actually the opening for their trunk. In
Norse mythology Norse, Nordic, or Scandinavian mythology is the body of myths belonging to the North Germanic peoples, stemming from Old Norse religion and continuing after the Christianization of Scandinavia, and into the Nordic folklore of the modern per ...
, echinoderm shells (the round five-part button left over from a sea urchin) were associated with the god
Thor Thor (; from non, Þórr ) is a prominent god in Germanic paganism. In Norse mythology, he is a hammer-wielding god associated with lightning, thunder, storms, sacred groves and trees, strength, the protection of humankind, hallowing, ...
, 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 Suffolk () is a ceremonial county of England in East Anglia. It borders Norfolk to the north, Cambridgeshire to the west and Essex to the south; the North Sea lies to the east. The county town is Ipswich; other important towns include ...
, a different species was used as a good-luck charm by bakers, who referred to them as fairy loaves, associating them with the similarly shaped loaves of bread they baked.


Early modern explanations

More scientific views of fossils emerged during the
Renaissance The Renaissance ( , ) , from , with the same meanings. is a period in European history marking the transition from the Middle Ages to modernity and covering the 15th and 16th centuries, characterized by an effort to revive and surpass ide ...
.
Leonardo da Vinci Leonardo di ser Piero da Vinci (15 April 14522 May 1519) was an Italian polymath of the High Renaissance who was active as a painter, draughtsman, engineer, scientist, theorist, sculptor, and architect. While his fame initially rested on ...
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 Robert Hooke FRS (; 18 July 16353 March 1703) was an English polymath active as a scientist, natural philosopher and architect, who is credited to be one of two scientists to discover microorganisms in 1665 using a compound microscope that ...
(1635–1703) included
micrograph A micrograph or photomicrograph is a photograph or digital image taken through a microscope or similar device to show a magnified image of an object. This is opposed to a macrograph or photomacrograph, an image which is also taken on a m ...
s of fossils in his '' Micrographia'' 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 (1769–1839), an English canal engineer, observed that rocks of different ages (based on the
law of superposition The law of superposition is an axiom that forms one of the bases of the sciences of geology, archaeology, and other fields pertaining to geological stratigraphy. In its plainest form, it states that in undeformed stratigraphic sequences, the ...
) preserved different assemblages of fossils, and that these assemblages succeeded one another in a regular and determinable order. He observed that rocks from distant locations could be correlated based on the fossils they contained. He termed this the principle of ''faunal succession''. This principle became one of Darwin's chief pieces of evidence that biological evolution was real.
Georges Cuvier Jean Léopold Nicolas Frédéric, Baron Cuvier (; 23 August 1769 – 13 May 1832), known as Georges Cuvier, was a French naturalist and zoologist, sometimes referred to as the "founding father of paleontology". Cuvier was a major figure in na ...
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 Ichthyosaurs (Ancient Greek for "fish lizard" – and ) are large extinct marine reptiles. Ichthyosaurs belong to the order known as Ichthyosauria or Ichthyopterygia ('fish flippers' – a designation introduced by Sir Richard Owen in 1842, alt ...
and a complete plesiosaurus 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 Phyla, the plural of ''phylum'', may refer to: * Phylum, a biological taxon between Kingdom and Class * by analogy, in linguistics, a large division of possibly related languages, or a major language family which is not subordinate to another Phy ...
) 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. 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
eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacter ...
s. 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 ...
or determining the age of rocks based on embedded fossils. For the first 150 years of
geology Geology () is a branch of natural science concerned with Earth and other Astronomical object, astronomical objects, the features or rock (geology), rocks of which it is composed, and the processes by which they change over time. Modern geology ...
, biostratigraphy and superposition were the only means for determining the relative age of rocks. The
geologic time scale The geologic time scale, or geological time scale, (GTS) is a representation of time based on the rock record of Earth. It is a system of chronological dating that uses chronostratigraphy (the process of relating strata to time) and geochr ...
was developed based on the relative ages of rock strata as determined by the early paleontologists and stratigraphers. Since the early years of the twentieth century,
absolute dating Absolute dating is the process of determining an age on a specified chronology in archaeology and geology. Some scientists prefer the terms chronometric or calendar dating, as use of the word "absolute" implies an unwarranted certainty of accurac ...
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, 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 Evolutionary biology is the subfield of biology that studies the evolutionary processes (natural selection, common descent, speciation) that produced the diversity of life on Earth. It is also defined as the study of the history of life ...
to share the interdisciplinary task of outlining the tree of life, which inevitably leads backwards in time to Precambrian microscopic life when cell structure and functions evolved. Earth's deep time in the Proterozoic and deeper still in the Archean is only "recounted by microscopic fossils and subtle chemical signals." Molecular biologists, using
phylogenetics In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups ...
, can compare protein
amino acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha ...
or
nucleotide Nucleotides are organic molecules consisting of a nucleoside and a phosphate. They serve as monomeric units of the nucleic acid polymers – deoxyribonucleic acid (DNA) and ribonucleic acid (RNA), both of which are essential biomolecu ...
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 Trilobites (; meaning "three lobes") are extinct marine arthropods that form the class Trilobita. Trilobites form one of the earliest-known groups of arthropods. The first appearance of trilobites in the fossil record defines the base of the ...
genus supported the hypothesis that modifications to the arrangement of the trilobite's eye lenses proceeded by fits and starts over millions of years during the
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, wh ...
. 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 Stephen Jay Gould (; September 10, 1941 – May 20, 2002) was an American paleontologist, evolutionary biologist, and historian of science. He was one of the most influential and widely read authors of popular science of his generation. Goul ...
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 A synchrotron is a particular type of cyclic particle accelerator, descended from the cyclotron, in which the accelerating particle beam travels around a fixed closed-loop path. The magnetic field which bends the particle beam into its closed ...
X-ray An X-ray, or, much less commonly, X-radiation, is a penetrating form of high-energy electromagnetic radiation. Most X-rays have a wavelength ranging from 10 picometers to 10  nanometers, corresponding to frequencies in the range 30&nb ...
tomographic analysis of early Cambrian bilaterian
embryo An embryo is an initial stage of development of a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male spe ...
nic 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'' 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 A germ layer is a primary layer of cells that forms during embryonic development. The three germ layers in vertebrates are particularly pronounced; however, all eumetazoans (animals that are sister taxa to the sponges) produce two or three p ...
embryonic development. These 543-million-year-old embryos support the emergence of some aspects of
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
development earlier than previously thought in the late Proterozoic. The preserved embryos from
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's List of countries and dependencies by population, most populous country, with a Population of China, population exceeding 1.4 billion, slig ...
and
Siberia Siberia ( ; rus, Сибирь, r=Sibir', p=sʲɪˈbʲirʲ, a=Ru-Сибирь.ogg) is an extensive geographical region, constituting all of North Asia, from the Ural Mountains in the west to the Pacific Ocean in the east. It has been a part ...
underwent rapid 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, 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 chiti ...
a. Despite significant advances in uncovering and identifying paleontological specimens, it is generally accepted that the fossil record is vastly incomplete. Approaches for measuring the completeness of the fossil record have been developed for numerous subsets of species, including those grouped taxonomically, temporally, environmentally/geographically, or in sum. This encompasses the subfield of taphonomy 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 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 A placebo ( ) is a substance or treatment which is designed to have no therapeutic value. Common placebos include inert tablets (like sugar pills), inert injections (like Saline (medicine), saline), sham surgery, and other procedures. In general ...
, though fossil material might conceivably have some antacid activity or supply some essential minerals. The use of dinosaur bones as "dragon bones" has persisted in
Traditional Chinese medicine Traditional Chinese medicine (TCM) is an alternative medical practice drawn from traditional medicine in China. It has been described as "fraught with pseudoscience", with the majority of its treatments having no logical mechanism of acti ...
into modern times, with mid-Cretaceous dinosaur bones being used for the purpose in
Ruyang County Ruyang County () is a county in the west of Henan province, China, under the jurisdiction of the prefecture-level city of Luoyang. Ruyang County has been the site of several major discoveries of fossilized dinosaur 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, 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 cuttlefis ...
fossils, each approximately 1.5 cm across File:Priscacara liops Green River Formation.jpg,
Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', ...
fossil fish ''Priscacara liops'' from the
Green River Formation The Green River Formation is an Eocene geologic formation that records the sedimentation in a group of intermountain lakes in three basins along the present-day Green River in Colorado, Wyoming, and Utah. The sediments are deposited in very f ...
of Wyoming File:Trilobite2.jpg, A permineralized
trilobite Trilobites (; meaning "three lobes") are extinct marine arthropods that form the class Trilobita. Trilobites form one of the earliest-known groups of arthropods. The first appearance of trilobites in the fossil record defines the base of the ...
, '' Asaphus kowalewskii'' 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'' teeth. The latter was found in the Sahara Desert. File:The fossils from Cretaceous age found in Lebanon.jpg, Fossil
shrimp Shrimp are crustaceans (a form of shellfish) with elongated bodies and a primarily swimming mode of locomotion – most commonly Caridea and Dendrobranchiata of the decapod order, although some crustaceans outside of this order are ref ...
(
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 Petrified Forest National Park is an American national park in Navajo County, Arizona, Navajo and Apache County, Arizona, Apache counties in northeastern Arizona. Named for its large deposits of petrified wood, the park covers about , encompassin ...
, Arizona File:Petrified Araucaria cone from patagonia-Edit3.jpg, Petrified cone 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 Patagonia () refers to a geographical region that encompasses the southern end of South America, governed by Argentina and Chile. The region comprises the southern section of the Andes Mountains with lakes, fjords, temperate rainforests, and g ...
,
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the List of South American countries by area, second-largest ...
dating from the Jurassic Period (approx. 210 Ma) File:CyprusPlioceneGastropod.JPG, A fossil
gastropod The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. T ...
from the
Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58Cyprus Cyprus ; tr, Kıbrıs (), officially the Republic of Cyprus,, , lit: Republic of Cyprus is an island country located south of the Anatolian Peninsula in the eastern Mediterranean Sea. Its continental position is disputed; while it is ...
. A serpulid worm is attached. File:OrhtocerasNautiloid092313.jpg,
Silurian The Silurian ( ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the shortest period of the Paleoz ...
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 ''Micraster'' is an extinct genus of echinoids from the Late Cretaceous to the early Eocene. Its remains have been found in Africa, Antarctica, Europe, and North America. Micraster was an infaunal echinoid living in a burrow below the sedi ...
'' echinoid fossil from England File:Productid Permian Texas.JPG, Productid brachiopod ventral valve; Roadian,
Guadalupian The Guadalupian is the second and middle series/ epoch of the Permian. The Guadalupian was preceded by the Cisuralian and followed by the Lopingian. It is named after the Guadalupe Mountains of New Mexico and Texas, and dates between 272.95 � ...
(Middle
Permian The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleo ...
); Glass Mountains, Texas. File:Fossil agatized coral Florida.JPG, Agatized
coral Corals are marine invertebrates within the class Anthozoa of the phylum Cnidaria. They typically form compact colonies of many identical individual polyps. Coral species include the important reef builders that inhabit tropical oceans and se ...
from the Hawthorn Group (
Oligocene The Oligocene ( ) is a geologic epoch of the Paleogene Period and extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that define the epoch are well identified but t ...
Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...
),
Florida Florida is a state located in the Southeastern region of the United States. Florida is bordered to the west by the Gulf of Mexico, to the northwest by Alabama, to the north by Georgia, to the east by the Bahamas and Atlantic Ocean, and ...
. 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 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

*
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,