A fossil (from Classical Latin fossilis; literally, "obtained by digging")[1] is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in amber, hair, petrified wood, oil, coal, and DNA remnants. The totality of fossils is known as the fossil record.

Paleontology is the study of fossils: their age, method of formation, and evolutionary significance. Specimens are usually considered to be fossils if they are over 10,000 years old.[2] The oldest fossils are around 3.48 billion years old[3][4][5] to 4.1 billion years old.[6][7] The observation in the 19th century that certain fossils were associated with certain rock strata led to the recognition of a geological timescale and the relative ages of different fossils. The development of radiometric dating techniques in the early 20th century allowed scientists to quantitatively measure the absolute ages of rocks and the fossils they host.

There are many processes that lead to fossilization, including permineralization, casts and molds, authigenic mineralization, replacement and recrystallization, adpression, carbonization, and bioimmuration.

Seymouria Fossil
Fossil of a Seymouria (extinct)

Fossils vary in size from one-micrometre (1 µm) bacteria[8] to dinosaurs and trees, many meters long and weighing many tons. A fossil normally preserves only a portion of the deceased organism, usually that portion that was partially mineralized during life, such as the bones and teeth of vertebrates, or the chitinous or calcareous exoskeletons of invertebrates. Fossils may also consist of the marks left behind by the organism while it was alive, such as animal tracks or feces (coprolites). These types of fossil are called trace fossils or ichnofossils, as opposed to body fossils. Some fossils are biochemical and are called chemofossils or biosignatures.

Fossilization processes

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


Permian Silicified Sclerobionts
Silicified (replaced with silica) fossils from the Road Canyon Formation (Middle Permian of Texas)

Permineralization is a process of fossilization that occurs when an organism is buried. The empty spaces within an organism (spaces filled with liquid or gas during life) become filled with mineral-rich groundwater. Minerals precipitate from the groundwater, occupying the empty spaces. This process can occur in very small spaces, such as within the cell wall of a plant cell. Small scale permineralization can produce very detailed fossils. For permineralization to occur, the organism must become covered by sediment soon after death or soon after the initial decay process. The degree to which the remains are decayed when covered determines the later details of the fossil. Some fossils consist only of skeletal remains or teeth; other fossils contain traces of skin, feathers or even soft tissues. This is a form of diagenesis.

Casts and molds

Aviculopecten subcardiformis01
External mold of a bivalve from the Logan Formation, Lower Carboniferous, Ohio

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 hole is later filled with other minerals, it is a cast. An endocast or internal mold is formed when sediments or minerals fill the internal cavity of an organism, such as the inside of a bivalve or snail or the hollow of a skull.

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

Recrystallized scleractinian coral (aragonite to calcite) from the Jurassic of southern Israel

Replacement occurs when the shell, bone or other tissue is replaced with another mineral. In some cases mineral replacement of the original shell occurs so gradually and at such fine scales that microstructural features are preserved despite the total loss of original material. A shell is said to be recrystallized when the original skeletal compounds are still present but in a different crystal form, as from aragonite to calcite.

Adpression (compression-impression)

Compression fossils, such as those of fossil ferns, are the result of chemical reduction of the complex organic molecules composing the organism's tissues. In this case the fossil consists of original material, albeit in a geochemically altered state. This chemical change is an expression of diagenesis. Often what remains is a carbonaceous film known as a phytoleim, in which case the fossil is known as a compression. Often, however, the phytoleim is lost and all that remains is an impression of the organism in the rock—an impression fossil. In many cases, however, compressions and impressions occur together. For instance, when the rock is broken open, the phytoleim will often be attached to one part (compression), whereas the counterpart will just be an impression. For this reason, one term covers the two modes of preservation: adpression.[9]

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.[10][11][12][13] 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.[14] However, a slightly older study based on eight taxa ranging in time from the Devonian to the Jurassic found that reasonably well-preserved fibrils that probably represent collagen were preserved in all these fossils, and that the quality of preservation depended mostly on the arrangement of the collagen fibers, with tight packing favoring good preservation.[15] There seemed to be no correlation between geological age and quality of preservation, within that timeframe.


Carbonaceous films are thin coatings which consist predominantly of the chemical element carbon. The soft tissues of organisms are made largely of organic carbon compounds and during diagenesis under reducing conditions only a thin film of carbon residue is left which forms a silhouette of the original organism.


The star-shaped holes (Catellocaula vallata) in this Upper Ordovician bryozoan represent a soft-bodied organism preserved by bioimmuration in the bryozoan skeleton.[16]

Bioimmuration occurs when a skeletal organism overgrows or otherwise subsumes another organism, preserving the latter, or an impression of it, within the skeleton.[17] Usually it is a sessile skeletal organism, such as a bryozoan or an oyster, which grows along a substrate, covering other sessile sclerobionts. Sometimes the bioimmured organism is soft-bodied and is then preserved in negative relief as a kind of external mold. There are also cases where an organism settles on top of a living skeletal organism that grows upwards, preserving the settler in its skeleton. Bioimmuration is known in the fossil record from the Ordovician[18] to the Recent.[17]


Estimating dates

Paleontology seeks to map out how life evolved across geologic time. A substantial hurdle is the difficulty of working out fossil ages. Beds that preserve fossils typically lack the radioactive elements needed for radiometric dating. This technique is our only means of giving rocks greater than about 50 million years old an absolute age, and can be accurate to within 0.5% or better.[19] 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.[19]


Consequently, palaeontologists rely on stratigraphy to date fossils. Stratigraphy is the science of deciphering the "layer-cake" that is the sedimentary record.[20] 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.[21] Because rock sequences are not continuous, but may be broken up by faults or periods of erosion, it is very difficult to match up rock beds that are not directly adjacent. However, fossils of species that survived for a relatively short time can be used to match isolated rocks: this technique is called biostratigraphy. For instance, the conodont Eoplacognathus pseudoplanus has a short range in the Middle Ordovician period.[22] 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.[23] 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 continents.[23] Family-tree relationships also help to narrow down the date when lineages first appeared. For instance, if fossils of B or C date to X million years ago and the calculated "family tree" says A was an ancestor of B and C, then A must have evolved earlier.

It is also possible to estimate how long ago two living clades diverged, in other words approximately how long ago their last common ancestor must have lived, by assuming that DNA mutations accumulate at a constant rate. These "molecular clocks", however, are fallible, and provide only approximate timing: for example, they are not sufficiently precise and reliable for estimating when the groups that feature in the Cambrian explosion first evolved,[24] and estimates produced by different techniques may vary by a factor of two.[25]


Fossil record gaps 20131028.pdf
Some of the most remarkable gaps in the fossil record (as of October 2013) show slanting toward organisms with hard parts.

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.[26] 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.[27]

The fossil record is strongly biased toward organisms with hard-parts, leaving most groups of soft-bodied organisms with little to no role.[26] It is replete with the mollusks, the vertebrates, the echinoderms, the brachiopods and some groups of arthropods.[28]



Fossil sites with exceptional preservation—sometimes including preserved soft tissues—are known as Lagerstätten - German for "storage places". These formations may have resulted from carcass burial in an anoxic environment with minimal bacteria, thus slowing decomposition. Lagerstätten span geological time from the Cambrian period to the present. Worldwide, some of the best examples of near-perfect fossilization are the Cambrian Maotianshan shales and Burgess Shale, the Devonian Hunsrück Slates, the Jurassic Solnhofen limestone, and the Carboniferous Mazon Creek localities.


Stromatolites Cochabamba
Lower Proterozoic stromatolites from Bolivia, South America

Stromatolites are layered accretionary structures formed in shallow water by the trapping, binding and cementation of sedimentary grains by biofilms of microorganisms, especially cyanobacteria.[29] Stromatolites provide some of the most ancient fossil records of life on Earth, dating back more than 3.5 billion years ago.[30]

Stromatolites were much more abundant in Precambrian times. While older, Archean fossil remains are presumed to be colonies of cyanobacteria, younger (that is, Proterozoic) fossils may be primordial forms of the eukaryote chlorophytes (that is, green algae). One genus of stromatolite very common in the geologic record is Collenia. The earliest stromatolite of confirmed microbial origin dates to 2.724 billion years ago.[31]

A 2009 discovery provides strong evidence of microbial stromatolites extending as far back as 3.45 billion years ago.[32]

Stromatolites are a major constituent of the fossil record for life's first 3.5 billion years, peaking about 1.25 billion years ago.[32] They subsequently declined in abundance and diversity,[33] 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.[34][35][36]

The connection between grazer and stromatolite abundance is well documented in the younger Ordovician 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.[37] 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.[38]

While prokaryotic cyanobacteria themselves reproduce asexually through cell division, they were instrumental in priming the environment for the evolutionary development of more complex eukaryotic organisms. Cyanobacteria (as well as extremophile Gammaproteobacteria) are thought to be largely responsible for increasing the amount of oxygen in the primeval earth's atmosphere through their continuing photosynthesis. Cyanobacteria use water, carbon dioxide and sunlight to create their food. A layer of mucus often forms over mats of cyanobacterial cells. In modern microbial mats, debris from the surrounding habitat can become trapped within the mucus, which can be cemented by the calcium carbonate to grow thin laminations of limestone. These laminations can accrete over time, resulting in the banded pattern common to stromatolites. The domal morphology of biological stromatolites is the result of the vertical growth necessary for the continued infiltration of sunlight to the organisms for photosynthesis. Layered spherical growth structures termed 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.[31]

The Zebra River Canyon area of the Kubis platform in the deeply dissected Zaris Mountains of southwestern Namibia provides an extremely well exposed example of the thrombolite-stromatolite-metazoan reefs that developed during the Proterozoic period, the stromatolites here being better developed in updip locations under conditions of higher current velocities and greater sediment influx.[39]



Index fossils
Examples of index fossils

Index fossils (also known as guide fossils, indicator fossils or zone fossils) are fossils used to define and identify geologic periods (or faunal stages). They work on the premise that, although different sediments may look different depending on the conditions under which they were deposited, they may include the remains of the same species 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.


A coprolite of a carnivorous dinosaur found in southwestern Saskatchewan

Trace fossils consist mainly of tracks and burrows, but also include coprolites (fossil feces) and marks left by feeding.[40][41] 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.[42] 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 earthworms).[41]

Coprolites are classified as trace fossils as opposed to body fossils, as they give evidence for the animal's behaviour (in this case, diet) rather than morphology. They were first described by William Buckland in 1829. Prior to this they were known as "fossil fir cones" and "bezoar stones." They serve a valuable purpose in paleontology because they provide direct evidence of the predation and diet of extinct organisms.[43] Coprolites may range in size from a few millimetres to over 60 centimetres.


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.[44] 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 about 1 mm

Microfossil is a descriptive term applied to fossilized plants and animals whose size is just at or below the level at which the fossil can be analyzed by the naked eye. A commonly applied cutoff point between "micro" and "macro" fossils is 1 mm. Microfossils may either be complete (or near-complete) organisms in themselves (such as the marine plankters foraminifera and coccolithophores) or component parts (such as small teeth or spores) of larger animals or plants. Microfossils are of critical importance as a reservoir of paleoclimate information, and are also commonly used by biostratigraphers to assist in the correlation of rock units.


Leptofoenus pittfieldae (male) rotated
Leptofoenus pittfieldae trapped in Dominican amber, from 20 to 16 million years ago

Fossil resin (colloquially called amber) is a natural polymer found in many types of strata throughout the world, even the Arctic. The oldest fossil resin dates to the Triassic, though most dates to the Cenozoic. The excretion of the resin by certain plants is thought to be an evolutionary adaptation for protection from insects and to seal wounds. Fossil resin often contains other fossils called inclusions that were captured by the sticky resin. These include bacteria, fungi, other plants, and animals. Animal inclusions are usually small invertebrates, predominantly arthropods such as insects and spiders, and only extremely rarely a vertebrate such as a small lizard. Preservation of inclusions can be exquisite, including small fragments of DNA.


Eroded Jurassic plesiosaur vertebral centrum found in the Lower Cretaceous Faringdon Sponge Gravels in Faringdon, England. An example of a remanié fossil.

A derived, reworked or remanié fossil is a fossil found in rock that accumulated significantly later than when the fossilized animal or plant died.[45] Reworked fossils are created by erosion exhuming (freeing) fossils from the rock formation in which they were originally deposited and their redeposition in an younger sedimentary deposit.


Petrified forest log 2 md
Petrified wood. The internal structure of the tree and bark are maintained in the permineralization process.
Petrified wood close 052615
Polished section of petrified wood showing annual rings

Fossil wood is wood that is preserved in the fossil record. Wood is usually the part of a plant that is best preserved (and most easily found). Fossil wood may or may not be petrified. The fossil wood may be the only part of the plant that has been preserved:[46] therefore such wood may get a special kind of botanical name. This will usually include "xylon" and a term indicating its presumed affinity, such as Araucarioxylon (wood of Araucaria or some related genus), Palmoxylon (wood of an indeterminate palm), or Castanoxylon (wood of an indeterminate chinkapin).[47]


Dodo-Skeleton Natural History Museum London England
A subfossil dodo skeleton

The term subfossil can be used to refer to remains, such as bones, nests, or defecations, 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.[48] The main importance of subfossil vs. fossil remains is that the former contain organic material, which can be used for radiocarbon dating or extraction and sequencing of DNA, protein, or other biomolecules. Additionally, isotope ratios can provide much information about the ecological conditions under which extinct animals lived. Subfossils are useful for studying the evolutionary history of an environment and can be important to studies in paleoclimatology.

Subfossils are often found in depositionary environments, such as lake sediments, oceanic sediments, and soils. Once deposited, physical and chemical weathering can alter the state of preservation.

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.[49] The oldest traces of life on Earth are fossils of this type, including carbon isotope anomalies found in zircons that imply the existence of life as early as 4.1 billion years ago.[6][7]


It has been suggested that biominerals could be important indicators of extraterrestrial life and thus could play an important role in the search for past or present life on the planet Mars. Furthermore, organic components (biosignatures) that are often associated with biominerals are believed to play crucial roles in both pre-biotic and biotic reactions.[50]

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 based on autotrophic, chemotrophic and/or chemolithoautotrophic microorganisms, as well as ancient water, including fluvio-lacustrine environments (plains related to ancient rivers or lakes) that may have been habitable.[51][52][53][54] The search for evidence of habitability, taphonomy (related to fossils), and organic carbon on the planet Mars is now a primary NASA objective.[51][52]


An example of a pseudofossil: Manganese dendrites on a limestone bedding plane from Solnhofen, Germany; scale in mm

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 dendrites, are formed by naturally occurring fissures in the rock that get filled up by percolating minerals. Other types of pseudofossils are kidney ore (round shapes in iron ore) and moss agates, which look like moss or plant leaves. Concretions, spherical or ovoid-shaped nodules found in some sedimentary strata, were once thought to be dinosaur eggs, and are often mistaken for fossils as well.

History of the study of fossils

Gathering fossils dates at least to the beginning of recorded history. The fossils themselves are referred to as the fossil record. The fossil record was one of the early sources of data underlying the study of evolution and continues to be relevant to the history of life on Earth. Paleontologists examine the fossil record to understand the process of evolution and the way particular species have evolved.

Ancient civilizations

Fossils have been visible and common throughout most of natural history, and so documented human interaction with them goes back as far as recorded history, or earlier.

There are many examples of paleolithic stone knives in Europe with fossil echinoderms set precisely at the hand grip, going all the way back to Homo heidelbergensis and neanderthals.[55] They also drilled holes through the center of these round shells and apparently used them as beads for necklaces.

The ancient Egyptians gathered fossils of species that resembled the bones of modern species they worshipped. The god Set was associated with the hippopotamus, therefore fossilized bones of hippo-like species were kept in that deity's temples.[56] Five-rayed fossil sea urchin shells were associated with the deity Sopdu, the Morning Star, equivalent of Venus in Roman mythology.[55]

Ceratopsian skulls are common in the Dzungarian Gate mountain pass in Asia, an area once famous for gold mines, as well as its endlessly cold winds. This has been attributed to legends of both gryphons and the land of Hyperborea

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, realized that fossil seashells from rocks were similar to those found on the beach, indicating the fossils were once living animals. Aristotle previously explained it in terms of vaporous exhalations,[57] which Avicenna modified into the theory of petrifying fluids (succus lapidificatus), later elaborated by Albert of Saxony in the 14th century and accepted in some form by most naturalists by the 16th century.[58]

Roman naturalist Pliny the Elder wrote of tongue stones, which he called glossopetra. These were shark teeth, thought by classical cultures to look like the tongues of people or snakes.[59] He also wrote about the Horn of Ammon, which are fossil ammonites, from which the species 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 ray-finned fish.[60]

The Plains tribes of North America are thought to have similarly associated fossils, such as the many intact pterosaur fossils naturally exposed in the region, with their own mythology of the thunderbird.[61]

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.[62]

In Japan, fossil shark teeth were associated with the mythical tengu, thought to be the razor-sharp claws of the creature, starting some time after the 8th century AD.[59]

In medieval China, the fossil bones of ancient mammals including Homo erectus were often mistaken for "dragon bones" and used as medicine and aphrodisiacs. In addition, some of these fossil bones are collected as "art" by scholars and they left scripts on it, indicating the time they got the collection. One good example is the famous scholar Huang Tingjian of the South Song Dynasty during the 11th century, who kept one seashell fossil with his poem engraved on it.[63] In the West fossilized sea creatures on mountainsides were seen as proof of the biblical deluge.

In 1027, the Persian Avicenna explained fossils' stoniness in The Book of Healing:

If what is said concerning the petrifaction of animals and plants is true, the cause of this (phenomenon) is a powerful mineralizing and petrifying virtue which arises in certain stony spots, or emanates suddenly from the earth during earthquake and subsidences, and petrifies whatever comes into contact with it. As a matter of fact, the petrifaction of the bodies of plants and animals is not more extraordinary than the transformation of waters.[64]

From the 13th century to the present day, scholars pointed out that the fossil skulls of Deinotherium giganteum, found in Crete and Greece, resemble cyclops skulls, and are quite possibly the origin of that Greek myth.[65][66]

Micraster coranguinum.4 - Cretacico superior
Fossil shells from the cretaceous era sea urchin, Micraster, were used in medieval times as both shepherd's crowns to protect houses, and as painted fairy loaves by bakers to bring luck to their bread-making.

In Norse mythology, echinoderm shells (the round five-part button left over from a sea urchin) were associated with the god Thor, not only being incorporated in thunderstones, representations of Thor's hammer and subsequent hammer-shaped crosses as Christianity was adopted, but also kept in houses to garner Thor's protection.[55]

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.[67] In Suffolk, a different species was used as a good-luck charm by bakers, who referred to them as fairy loaves, associating them with the similarly shaped loaves of bread they baked.[68][69]

Early modern explanations

More scientific views of fossils emerged during the Renaissance. Leonardo da Vinci concurred with Aristotle's view that fossils were the remains of ancient life.[70] For example, da Vinci noticed discrepancies with the biblical flood narrative as an explanation for fossil origins:

If the Deluge had carried the shells for distances of three and four hundred miles from the sea it would have carried them mixed with various other natural objects all heaped up together; but even at such distances from the sea we see the oysters all together and also the shellfish and the cuttlefish and all the other shells which congregate together, found all together dead; and the solitary shells are found apart from one another as we see them every day on the sea-shores.

And we find oysters together in very large families, among which some may be seen with their shells still joined together, indicating that they were left there by the sea and that they were still living when the strait of Gibraltar was cut through. In the mountains of Parma and Piacenza multitudes of shells and corals with holes may be seen still sticking to the rocks...."[71]

Rozprawa o přewratech kůry zemnj, Ichthyosaurus and Plesiosaurus
Ichthyosaurus and Plesiosaurus from the 1834 Czech edition of Cuvier's Discours sur les revolutions de la surface du globe

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.[59]

Robert Hooke (1635-1703) included micrographs of fossils in his Micrographia and was among the first to observe fossil forams. His observations on fossils, which he stated to be the petrified remains of creatures some of which no longer existed, were published posthumously in 1705.[72]

William Smith (1769–1839), an English canal engineer, observed that rocks of different ages (based on the law of superposition) preserved different assemblages of fossils, and that these assemblages succeeded one another in a regular and determinable order. He observed that rocks from distant locations could be correlated based on the fossils they contained. He termed this the principle of faunal succession. This principle became one of Darwin's chief pieces of evidence that biological evolution was real.

Georges Cuvier came to believe that most if not all the animal fossils he examined were remains of extinct species. This led Cuvier to become an active proponent of the geological school of thought called catastrophism. Near the end of his 1796 paper on living and fossil elephants he said:

All of these facts, consistent among themselves, and not opposed by any report, seem to me to prove the existence of a world previous to ours, destroyed by some kind of catastrophe.[73]

Interest in fossils, and geology more generally, expanded during the early nineteenth century. In Britain, Mary Anning's discoveries of fossils, including the first complete ichthyosaur and a complete plesiosaurus skeleton, sparked both public and scholarly interest.[74]

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 Period, now known to be about 540 million years old. He worried about the absence of older fossils because of the implications on the validity of his theories, but he expressed hope that such fossils would be found, noting that: "only a small portion of the world is known with accuracy." Darwin also pondered the sudden appearance of many groups (i.e. phyla) in the oldest known Cambrian fossiliferous strata.[75]

After Darwin

Since Darwin's time, the fossil record has been extended to between 2.3 and 3.5 billion years.[76] 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 eukaryotes.

The fossil record and faunal succession form the basis of the science of biostratigraphy or determining the age of rocks based on embedded fossils. For the first 150 years of geology, biostratigraphy and superposition were the only means for determining the relative age of rocks. The geologic time scale was developed based on the relative ages of rock strata as determined by the early paleontologists and stratigraphers.

Since the early years of the twentieth century, absolute dating methods, such as radiometric dating (including potassium/argon, argon/argon, uranium series, and, for very recent fossils, radiocarbon dating) have been used to verify the relative ages obtained by fossils and to provide absolute ages for many fossils. Radiometric dating has shown that the earliest known stromatolites are over 3.4 billion years old.

Modern era

Paleontology has joined with evolutionary biology to share the interdisciplinary task of outlining the tree of life, which inevitably leads backwards in time to Precambrian microscopic life when cell structure and functions evolved. Earth's deep time in the Proterozoic and deeper still in the Archean is only "recounted by microscopic fossils and subtle chemical signals."[78] Molecular biologists, using phylogenetics, can compare protein amino acid or nucleotide sequence homology (i.e., similarity) to evaluate taxonomy and evolutionary distances among organisms, with limited statistical confidence. The study of fossils, on the other hand, can more specifically pinpoint when and in what organism a mutation first appeared. Phylogenetics and paleontology work together in the clarification of science's still dim view of the appearance of life and its evolution.[79]

Phacopid trilobite Eldredgeops rana crassituberculata. The genus is named after Niles Eldredge.
Isocrinus nicoleti Encrinite Mt Carmel
Crinoid columnals (Isocrinus nicoleti) from the Middle Jurassic Carmel Formation at Mount Carmel Junction, Utah

Niles Eldredge's study of the Phacops trilobite genus supported the hypothesis that modifications to the arrangement of the trilobite's eye lenses proceeded by fits and starts over millions of years during the Devonian.[80] Eldredge's interpretation of the Phacops fossil record was that the aftermaths of the lens changes, but not the rapidly occurring evolutionary process, were fossilized. This and other data led Stephen Jay Gould and Niles Eldredge to publish their seminal paper on punctuated equilibrium in 1971.

Synchrotron X-ray tomographic analysis of early Cambrian bilaterian embryonic microfossils yielded new insights of metazoan evolution at its earliest stages. The tomography technique provides previously unattainable three-dimensional resolution at the limits of fossilization. Fossils of two enigmatic bilaterians, the worm-like Markuelia and a putative, primitive protostome, Pseudooides, provide a peek at germ layer embryonic development. These 543-million-year-old embryos support the emergence of some aspects of arthropod development earlier than previously thought in the late Proterozoic. The preserved embryos from China and Siberia 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 Arthropoda.[81]

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.[82] The problem is quite pronounced in China, where many specimens have been stolen.[83]

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.

Fossils as medicine

These is some medicinal and preventive use for some fossils. Largely the use of fossils as medicine is a matter of placebo effect. However, the consumption of certain fossils has been proven to help against stomach acidity and mineral depletion. The use of fossils to address health issues is rooted in traditional medicine and include the use of fossils as talismans. The specific fossil to use to alleviate or cure an illness is often based on its resemblance of the fossils and the symptoms or affected organ.[84]


Amonite Cropped

Three small ammonite fossils, each approximately 1.5 cm across

Priscacara liops Green River Formation

Eocene fossil fish Priscacara liops from the Green River Formation of Wyoming

Asaphus kowalewskii 3

A permineralized trilobite, Asaphus kowalewskii

Carcharodontosaurus and Megalodon teeth

Megalodon and Carcharodontosaurus teeth. The latter was found in the Sahara Desert.

The fossils from Cretaceous age found in Lebanon

Fossil shrimp (Cretaceous)


Petrified wood in Petrified Forest National Park, Arizona

Petrified Araucaria cone from patagonia-Edit3

Petrified cone of Araucaria mirabilis from Patagonia, Argentina dating from the Jurassic Period (approx. 210 Ma)


A fossil gastropod from the Pliocene of Cyprus. A serpulid worm is attached.

Eocene fossil flower, Clare Family Florissant Fossil Quarry, Florissant, Colorado, USA - 20100807

Eocene fossil flower from Florissant, Colorado


Micraster echinoid fossil from England

Productid Permian Texas

Productid brachiopod ventral valve; Roadian, Guadalupian (Middle Permian); Glass Mountains, Texas.

Fossil agatized coral Florida

Agatized coral from the Hawthorn Group (OligoceneMiocene), Florida. An example of preservation by replacement.

Fossils from Gotland beaches

Fossils from beaches of the Baltic Sea island of Gotland, placed on paper with 7 mm (0.28 inch) squares

See also


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Further reading

External links


Amber is fossilized tree resin, which 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 is used in jewelry. It has also been used as a healing agent in folk medicine.

There are five classes of amber, defined on the basis of their chemical constituents. Because it originates as a soft, sticky tree resin, amber sometimes contains animal and plant material as inclusions. Amber occurring in coal seams is also called resinite, and the term ambrite is applied to that found specifically within New Zealand coal seams.

Cradle of Humankind

The Cradle of Humankind is a paleoanthropological site about 50 km (31 mi) northwest of Johannesburg, South Africa, in the Gauteng province. Declared a World Heritage site by UNESCO in 1999, the site currently occupies 47,000 hectares (180 sq mi) and contains a complex of limestone caves. The registered name of the site in the list of World Heritage sites is Fossil Hominid Sites of South Africa.

The Sterkfontein Caves were the site of the discovery of a 2.3-million-year-old fossil Australopithecus africanus (nicknamed "Mrs. Ples"), found in 1947 by Robert Broom and John T. Robinson. The find helped corroborate the 1924 discovery of the juvenile Australopithecus africanus skull known as the "Taung Child", by Raymond Dart, at Taung in the North West Province of South Africa, where excavations still continue.

Nearby the site, but not in the site, the Rising Star Cave system contains the Dinaledi Chamber (chamber of stars), in which were discovered fifteen fossil skeletons of an extinct species of hominin, provisionally named Homo naledi.

Sterkfontein alone has produced more than a third of early hominid fossils ever found prior to 2010. The Dinaledi Chamber contains over 1,500 H. naledi fossils, the most extensive discovery of a single hominid species ever found in Africa.


Crustaceans (Crustacea ) form a large, diverse arthropod taxon which includes such familiar animals as crabs, lobsters, crayfish, shrimp, krill, woodlice, and barnacles.

The crustacean group is usually treated as a subphylum, and because of recent molecular studies it is now well accepted that the crustacean group is paraphyletic, and comprises all animals in the Pancrustacea clade other than hexapods. Some crustaceans are more closely related to insects and other hexapods than they are to certain other crustaceans.

The 67,000 described species range in size from Stygotantulus stocki at 0.1 mm (0.004 in), to the Japanese spider crab with a leg span of up to 3.8 m (12.5 ft) and a mass of 20 kg (44 lb). Like other arthropods, crustaceans have an exoskeleton, which they moult to grow. They are distinguished from other groups of arthropods, such as insects, myriapods and chelicerates, by the possession of biramous (two-parted) limbs, and by their larval forms, such as the nauplius stage of branchiopods and copepods.

Most crustaceans are free-living aquatic animals, but some are terrestrial (e.g. woodlice), some are parasitic (e.g. Rhizocephala, fish lice, tongue worms) and some are sessile (e.g. barnacles). The group has an extensive fossil record, reaching back to the Cambrian, and includes living fossils such as Triops cancriformis, which has existed apparently unchanged since the Triassic period. More than 10 million tons of crustaceans are produced by fishery or farming for human consumption, the majority of it being shrimp and prawns. Krill and copepods are not as widely fished, but may be the animals with the greatest biomass on the planet, and form a vital part of the food chain. The scientific study of crustaceans is known as carcinology (alternatively, malacostracology, crustaceology or crustalogy), and a scientist who works in carcinology is a carcinologist.


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, although the exact origin and timing of the evolution of dinosaurs is the subject of active research. They became the dominant terrestrial vertebrates after the Triassic–Jurassic extinction event 201 million years ago; their dominance continued through the Jurassic and Cretaceous periods. Reverse genetic engineering and the fossil record both demonstrate that birds are modern feathered dinosaurs, having evolved from earlier theropods during the late Jurassic Period. As such, birds were the only dinosaur lineage to survive the Cretaceous–Paleogene extinction event 66 million years ago. Dinosaurs can therefore be divided into avian dinosaurs, or birds; and non-avian dinosaurs, which are all dinosaurs other than birds. This article deals primarily with non-avian dinosaurs.

Dinosaurs are a varied group of animals from taxonomic, morphological and ecological standpoints. Birds, at over 10,000 living species, are the most diverse group of vertebrates besides perciform fish. Using fossil evidence, paleontologists have identified over 500 distinct genera and more than 1,000 different species of non-avian dinosaurs. Dinosaurs are represented on every continent by both extant species (birds) and fossil remains. Through the first half of the 20th century, before birds were recognized to be dinosaurs, most of the scientific community believed dinosaurs to have been sluggish and cold-blooded. Most research conducted since the 1970s, however, has indicated that all dinosaurs were active animals with elevated metabolisms and numerous adaptations for social interaction. Some were herbivorous, others carnivorous. Evidence suggests that egg-laying and nest-building are additional traits shared by all dinosaurs, avian and non-avian alike.

While dinosaurs were ancestrally bipedal, many extinct groups included quadrupedal species, and some were able to shift between these stances. Elaborate display structures such as horns or crests are common to all dinosaur groups, and some extinct groups developed skeletal modifications such as bony armor and spines. While the dinosaurs' modern-day surviving avian lineage (birds) are generally small due to the constraints of flight, many prehistoric dinosaurs (non-avian and avian) were large-bodied—the largest sauropod dinosaurs are estimated to have reached lengths of 39.7 meters (130 feet) and heights of 18 meters (59 feet) and were the largest land animals of all time. Still, the idea that non-avian dinosaurs were uniformly gigantic is a misconception based in part on preservation bias, as large, sturdy bones are more likely to last until they are fossilized. Many dinosaurs were quite small: Xixianykus, for example, was only about 50 cm (20 in) long.

Since the first dinosaur fossils were recognized in the early 19th century, mounted fossil dinosaur skeletons have been major attractions at museums around the world, and dinosaurs have become an enduring part of world culture. The large sizes of some dinosaur groups, as well as their seemingly monstrous and fantastic nature, have ensured dinosaurs' regular appearance in best-selling books and films, such as Jurassic Park. Persistent public enthusiasm for the animals has resulted in significant funding for dinosaur science, and new discoveries are regularly covered by the media.

Fossil fuel

A fossil fuel is a fuel formed by natural processes, such as anaerobic decomposition of buried dead organisms, containing energy originating in ancient photosynthesis.

The age of the organisms and their resulting fossil fuels is typically millions of years, and sometimes exceeds 650 million years.

Fossil fuels contain high percentages of carbon and include petroleum, coal, and natural gas.

Other commonly used derivatives include kerosene and propane.

Fossil fuels range from volatile materials with low carbon to hydrogen ratios like methane, to liquids like petroleum, to nonvolatile materials composed of almost pure carbon, like anthracite coal.

Methane can be found in hydrocarbon fields either alone, associated with oil, or in the form of methane clathrates.

The theory that fossil fuels formed from the fossilized remains of dead plants by exposure to heat and pressure in the Earth's crust over millions of years was first introduced by Andreas Libavius "in his 1597 Alchemia [Alchymia]" and later by Mikhail Lomonosov "as early as 1757 and certainly by 1763". The first use of the term "fossil fuel" was by the German chemist Caspar Neumann, in English translation in 1759.In 2017 the world's primary energy sources consisted of petroleum (34%), coal (28%), natural gas (23%), amounting to an 85% share for fossil fuels in primary energy consumption in the world.

Non-fossil sources in 2006 included nuclear (8.5%), hydroelectric (6.3%), and others (geothermal, solar, tidal, wind, wood, waste) amounting to 0.9%.

World energy consumption was growing at about 2.3% per year. In 2015 about 18% of worldwide consumption was from renewable sources.Although fossil fuels are continually being formed via natural processes, they are generally considered to be non-renewable resources because they take millions of years to form and the known viable reserves are being depleted much faster than new ones are being made.The use of fossil fuels raises serious environmental concerns.

The burning of fossil fuels produces around 21.3 billion tonnes (21.3 gigatonnes) of carbon dioxide (CO2) per year.

It is estimated that natural processes can only absorb about half of that amount, so there is a net increase of 10.65 billion tonnes of atmospheric carbon dioxide per year.

Carbon dioxide is a greenhouse gas that increases radiative forcing and contributes to global warming.

A global movement towards the generation of low-carbon renewable energy is underway to help reduce global greenhouse gas emissions.

Fossil fuel phase-out

Fossil fuel phase out refers to the discontinuation of the use of fossil fuels, through the decommissioning of operating fossil fuel-fired power plants, the prevention of the construction of new ones, and the use of alternative energy to replace the role of fossil fuels.

The purpose of fossil fuel phase-out is to reduce the negative externalities that use of fossil fuels cause. Negative externalities refer to the costs a certain activity has over people who did not choose to incur in them. A direct negative externality from fossil fuels' use is air pollution, and an indirect negative externality are mining accidents, that happen as a consequence of the extraction of fossil fuels. Fossil fuel burning contributes to climate change, as it releases greenhouse gas emissions.

Fossil fuel power station

A fossil fuel power station is a thermal power station which burns a fossil fuel such as coal, natural gas, or petroleum to produce electricity. Central station fossil fuel power plants are designed on a large scale for continuous operation. In many countries, such plants provide most of the electrical energy used. Fossil fuel power stations have machinery to convert the heat energy of combustion into mechanical energy, which then operates an electrical generator. The prime mover may be a steam turbine, a gas turbine or, in small plants, a reciprocating internal combustion engine. All plants use the energy extracted from expanding gas, either steam or combustion gases.

Although different energy conversion methods exist, all thermal power station conversion methods have efficiency limited by the Carnot efficiency and therefore produce waste heat.

By-products of fossil fuel power plant operation must be considered in their design and operation. The flue gas from combustion of the fossil fuels is discharged to the air. This gas contains carbon dioxide and water vapor, as well as other substances such as nitrogen oxides (NOx), sulfur oxides (SOx), mercury, traces of other metals, and, for coal-fired plants, fly ash. Solid waste ash from coal-fired boilers must also be removed. Some coal ash can be recycled for building materials.Fossil fueled power stations are major emitters of carbon dioxide (CO2), a greenhouse gas which is a major contributor to global warming.

The results of a recent study show that the net income available to shareholders of large companies could see a significant reduction from the greenhouse gas emissions liability related to only natural disasters in the United States from a single coal-fired power plant.

However, as of 2015, no such cases have awarded damages in the United States.

Per unit of electric energy, brown coal emits nearly two times as much CO2 as natural gas, and black coal emits somewhat less than brown.

Carbon capture and storage of emissions has been proposed to limit the environmental impact of fossil fuel power stations, but it is still at a demonstration stage.

Galaxy group

A galaxy group or group of galaxies (GrG) is an aggregation of galaxies comprising about 50 or fewer gravitationally bound members, each at least as luminous as the Milky Way (about 1010 times the luminosity of the Sun); collections of galaxies larger than groups that are first-order clustering are called galaxy clusters. The groups and clusters of galaxies can themselves be clustered, into superclusters of galaxies.

The Milky Way galaxy is part of a group of galaxies called the Local Group.


A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several such, but explicitly designated as the holotype. Under the International Code of Zoological Nomenclature (ICZN), a holotype is one of several kinds of name-bearing types. In the International Code of Nomenclature for algae, fungi, and plants (ICN) and ICZN the definitions of types are similar in intent but not identical in terminology or underlying concept.

For example, the holotype for the butterfly Lycaeides idas longinus is a preserved specimen of that species, held by the Museum of Comparative Zoology at Harvard University. An isotype is a duplicate of the holotype and is often made for plants, where holotype and isotypes are often pieces from the same individual plant or samples from the same gathering.

A holotype is not necessarily "typical" of that taxon, although ideally it should be. Sometimes just a fragment of an organism is the holotype, particularly in the case of a fossil. For example, the holotype of Pelorosaurus humerocristatus (Duriatitan), a large herbivorous dinosaur from the early Jurassic period, is a fossil leg bone stored at the Natural History Museum in London. Even if a better specimen is subsequently found, the holotype is not superseded.


The Hominidae (), whose members are known as great apes or hominids, are a taxonomic family of primates that includes eight extant species in four genera: Pongo, the Bornean, Sumatran and Tapanuli orangutan; Gorilla, the eastern and western gorilla; Pan, the common chimpanzee and the bonobo; and Homo, which includes modern humans and its extinct relatives (e.g., the Neanderthal), and ancestors, such as Homo erectus.Several revisions in classifying the great apes have caused the use of the term "hominid" to vary over time. Its original meaning referred only to humans (Homo) and their closest extinct relatives. That restrictive meaning has now been largely assumed by the term "hominin", which comprises all members of the human clade after the split from the chimpanzees (Pan). The current, 21st-century meaning of "hominid" includes all the great apes including humans. Usage still varies, however, and some scientists and laypersons still use "hominid" in the original restrictive sense; the scholarly literature generally shows the traditional usage until around the turn of the 21st century.Within the taxon Hominidae, a number of extant and known extinct, that is, fossil, genera are grouped with the humans, chimpanzees, and gorillas in the subfamily Homininae; others with orangutans in the subfamily Ponginae (see classification graphic below). The most recent common ancestor of all Hominidae lived roughly 14 million years ago, when the ancestors of the orangutans speciated from the ancestral line of the other three genera. Those ancestors of the family Hominidae had already speciated from the family Hylobatidae (the gibbons), perhaps 15 million to 20 million years ago.

Internal combustion engine

An internal combustion engine (ICE) is a heat engine where the combustion of a fuel occurs with an oxidizer (usually air) in a combustion chamber that is an integral part of the working fluid flow circuit. In an internal combustion engine, the expansion of the high-temperature and high-pressure gases produced by combustion applies direct force to some component of the engine. The force is applied typically to pistons, turbine blades, rotor or a nozzle. This force moves the component over a distance, transforming chemical energy into useful mechanical energy.

The first commercially successful internal combustion engine was created by Étienne Lenoir around 1859 and the first modern internal combustion engine was created in 1876 by Nikolaus Otto (see Otto engine).

The term internal combustion engine usually refers to an engine in which combustion is intermittent, such as the more familiar four-stroke and two-stroke piston engines, along with variants, such as the six-stroke piston engine and the Wankel rotary engine. A second class of internal combustion engines use continuous combustion: gas turbines, jet engines and most rocket engines, each of which are internal combustion engines on the same principle as previously described. Firearms are also a form of internal combustion engine.In contrast, in external combustion engines, such as steam or Stirling engines, energy is delivered to a working fluid not consisting of, mixed with, or contaminated by combustion products. Working fluids can be air, hot water, pressurized water or even liquid sodium, heated in a boiler. ICEs are usually powered by energy-dense fuels such as gasoline or diesel fuel, liquids derived from fossil fuels. While there are many stationary applications, most ICEs are used in mobile applications and are the dominant power supply for vehicles such as cars, aircraft, and boats.

Typically an ICE is fed with fossil fuels like natural gas or petroleum products such as gasoline, diesel fuel or fuel oil. There is a growing usage of renewable fuels like biodiesel for CI (compression ignition) engines and bioethanol or methanol for SI (spark ignition) engines. Hydrogen is sometimes used, and can be obtained from either fossil fuels or renewable energy.

Lucy (Australopithecus)

Lucy is the common name of AL 288-1, several hundred pieces of bone fossils representing 40 percent of the skeleton of a female of the hominin species Australopithecus afarensis. In Ethiopia, the assembly is also known as Dinkinesh, which means "you are marvelous" in the Amharic language. Lucy was discovered in 1974 in Africa, near the village Hadar in the Awash Valley of the Afar Triangle in Ethiopia, by paleoanthropologist Donald Johanson of the Cleveland Museum of Natural History.The Lucy specimen is an early australopithecine and is dated to about 3.2 million years ago. The skeleton presents a small skull akin to that of non-hominin apes, plus evidence of a walking-gait that was bipedal and upright, akin to that of humans (and other hominins); this combination supports the view of human evolution that bipedalism preceded increase in brain size. A 2016 study proposes that Australopithecus afarensis was also to a large extent tree-dwelling, though the extent of this is debated."Lucy" acquired her name from the song "Lucy in the Sky with Diamonds" by The Beatles, which was played loudly and repeatedly in the expedition camp all evening after the excavation team's first day of work on the recovery site. After public announcement of the discovery, Lucy captured much public interest, becoming a household name at the time.

Lucy became famous worldwide, and the story of her discovery and reconstruction was published in a book by Johanson. Beginning in 2007, the fossil assembly and associated artifacts were exhibited publicly in an extended six-year tour of the United States; the exhibition was called Lucy's Legacy: The Hidden Treasures of Ethiopia. There was discussion of the risks of damage to the unique fossils, and other museums preferred to display casts of the fossil assembly. The original fossils were returned to Ethiopia in 2013, and subsequent exhibitions have used casts.


Megalodon (Carcharocles megalodon), meaning "big tooth", is an extinct species of shark that lived approximately 23 to 2.6 million years ago (mya), during the Early Miocene to the end of the Pliocene. It was formerly thought to be a member of the Lamnidae family, making it closely related to the great white shark (Carcharodon carcharias). However presently there is near unanimous consensus that it belongs to the extinct family Otodontidae, which diverged from the ancestry of the great white shark during the Early Cretaceous. Its genus placement is still debated, authors placing it in either Carcharocles, Megaselachus, Otodus, or Procarcharodon.

Scientists suggest that megalodon looked like a stockier version of the great white shark, though it may have looked similar to the basking shark (Cetorhinus maximus) or the sand tiger shark (Carcharias taurus). Regarded as one of the largest and most powerful predators to have ever lived, fossil remains of megalodon suggest that this giant shark reached a maximum length of 18 meters (59 ft) with the average size being 10.5 meters (34 ft). Their large jaws could exert a bite force of up to 110,000 to 180,000 newtons (25,000 to 40,000 lbf). Their teeth were thick and robust, built for grabbing prey and breaking bone.

Megalodon probably had a major impact on the structure of marine communities. The fossil record indicates that it had a cosmopolitan distribution. It probably targeted large prey, such as whales, seals, and sea turtles. Juveniles inhabited warm coastal waters and fed on fish and small whales. Unlike the great white, which attacks prey from the soft underside, megalodon probably used its strong jaws to break through the chest cavity and puncture the heart and lungs of its prey.

The animal faced competition from whale-eating cetaceans, such as Livyatan and other macroraptorial sperm whales, and smaller ancestral killer whales such as Orcinus citoniensis. As the shark preferred warmer waters, it is thought that oceanic cooling associated with the onset of the ice ages, coupled with the lowering of sea levels and resulting loss of suitable nursery areas, may have also contributed to its decline. A reduction in the diversity of baleen whales and a shift in their distribution toward polar regions may have reduced megalodon's primary food source. The extinction of the shark appeared to affect other animals; for example, the size of baleen whales increased significantly after the shark had disappeared.


Monotremes (from Greek μονός, monos ("single") and τρῆμα, trema ("hole"), referring to the cloaca) are one of the three main groups of living mammals, along with placentals (Eutheria) and marsupials (Metatheria). The monotremes are typified by structural differences in their brains, jaws, digestive tract, reproductive tract, and other body parts compared to the more common mammalian types. In addition they lay eggs rather than bear live young, but, like marsupials, they store their newly hatched, larvae-like, developing puggles in a pouch, and, like all mammals, the female monotremes nurse their young with milk.

Monotremes are traditionally referred to as the mammalian subclass Prototheria. The only surviving examples of monotremes are all indigenous to Australia and New Guinea although there is evidence that they were once more widespread including some extinct species in South America. The existing monotreme species are the platypus and four species of echidnas. There is currently some debate regarding monotreme taxonomy.

Non-renewable resource

A non-renewable resource (also called a finite resource) is a resource of economic value that cannot be readily replaced by natural means at a quick enough pace to keep up with consumption. An example is carbon-based fossil fuel. The original organic material, with the aid of heat and pressure, becomes a fuel such as oil or gas. Earth minerals and metal ores, fossil fuels (coal, petroleum, natural gas) and groundwater in certain aquifers are all considered non-renewable resources, though individual elements are always conserved (except in nuclear reactions).

On the other hand, resources such as timber (when harvested sustainably) and wind (used to power energy conversion systems) are considered renewable resources, largely because their localized replenishment can occur within time frames meaningful to humans too.


Paleobotany, also spelled as palaeobotany (from the Greek words paleon = old and "botany", study of plants), is the branch of paleontology or paleobiology dealing with the recovery and identification of plant remains from geological contexts, and their use for the biological reconstruction of past environments (paleogeography), and both the evolutionary history of plants, with a bearing upon the evolution of life in general. A synonym is paleophytology. Paleobotany includes the study of terrestrial plant fossils, as well as the study of prehistoric marine photoautotrophs, such as photosynthetic algae, seaweeds or kelp. A closely related field is palynology, which is the study of fossilized and extant spores and pollen.

Paleobotany is important in the reconstruction of ancient ecological systems and climate, known as paleoecology and paleoclimatology respectively; and is fundamental to the study of green plant development and evolution. Paleobotany has also become important to the field of archaeology, primarily for the use of phytoliths in relative dating and in paleoethnobotany.

The emergence of paleobotany as a scientific discipline can be seen in the early 19th century, especially in the works of the German palaeontologist Ernst Friedrich von Schlotheim, the Czech (Bohemian) nobleman and scholar Kaspar Maria von Sternberg, and the French botanist Adolphe-Théodore Brongniart.


Paleontology or palaeontology () 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 fossils to determine organisms' evolution and interactions with each other and their environments (their paleoecology). Paleontological observations have been documented as far back as the 5th century BC. The science became established in the 18th century as a result of Georges Cuvier's work on comparative anatomy, and developed rapidly in the 19th century. The term itself originates from Greek παλαιός, palaios, "old, ancient", ὄν, on (gen. ontos), "being, creature" and λόγος, logos, "speech, thought, study".Paleontology lies on the border between biology and geology, but differs from archaeology in that it excludes the study of anatomically modern humans. It now uses techniques drawn from a wide range of sciences, including biochemistry, mathematics, and engineering. Use of all these techniques has enabled paleontologists to discover much of the evolutionary history of life, almost all the way back to when Earth became capable of supporting life, about 3.8 billion years ago. As knowledge has increased, paleontology has developed specialised sub-divisions, some of which focus on different types of fossil organisms while others study ecology and environmental history, such as ancient climates.

Body fossils and trace fossils are the principal types of evidence about ancient life, and geochemical evidence has helped to decipher the evolution of life before there were organisms large enough to leave body fossils. Estimating the dates of these remains is essential but difficult: sometimes adjacent rock layers allow radiometric dating, which provides absolute dates that are accurate to within 0.5%, but more often paleontologists have to rely on relative dating by solving the "jigsaw puzzles" of biostratigraphy. Classifying ancient organisms is also difficult, as many do not fit well into the Linnaean taxonomy that is commonly used for classifying living organisms, and paleontologists more often use cladistics to draw up evolutionary "family trees". The final quarter of the 20th century saw the development of molecular phylogenetics, which investigates how closely organisms are related by measuring how similar the DNA is in their genomes. Molecular phylogenetics has also been used to estimate the dates when species diverged, but there is controversy about the reliability of the molecular clock on which such estimates depend.


Sauropsida ("lizard faces") is a taxonomic clade that includes both reptiles and birds. All Tetrapoda except Amphibia are Amniota, and all Amniota except Synapsida, including Mammalia, are Sauropsida. This clade includes Parareptilia and other extinct clades. All living sauropsids are members of the subgroup Diapsida, the Parareptilia having died out 200 million years ago. The term originated in 1864 with Thomas Henry Huxley, who grouped birds with reptiles based on fossil evidence.

Trace fossil

A trace fossil, also ichnofossil ( ; from Greek: ἴχνος ikhnos "trace, track"), is a geological record of biological activity. Ichnology is the study of such traces, and is the work of ichnologists. Trace fossils may consist of impressions made on or in the substrate by an organism: for example, burrows, borings (bioerosion), urolites (erosion caused by evacuation of liquid wastes), footprints and feeding marks, and root cavities. The term in its broadest sense also includes the remains of other organic material produced by an organism — for example coprolites (fossilized droppings) or chemical markers — or sedimentological structures produced by biological means - for example, stromatolites. Trace fossils contrast with body fossils, which are the fossilized remains of parts of organisms' bodies, usually altered by later chemical activity or mineralization.

Sedimentary structures, for example those produced by empty shells rolling along the sea floor, are not produced through the behaviour of an organism and not considered trace fossils.

The study of traces - ichnology - divides into paleoichnology, or the study of trace fossils, and neoichnology, the study of modern traces. Ichnological science offers many challenges, as most traces reflect the behaviour — not the biological affinity — of their makers. Accordingly, researchers classify trace fossils into form genera, based on their appearance and on the implied behaviour, or ethology, of their makers.

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