Fossils (from Latin fossus, literally “having been dug up”) are the preserved remains or traces of animals (also known as zoolites), plants, and other organisms from the remote past. The totality of fossils, both discovered and undiscovered, and their placement in fossiliferous (fossil-containing) rock formations andsedimentary layers (strata) is known as the fossil record.

The study of fossils across geological time, how they were formed, and the evolutionary relationships between taxa (phylogeny) are some of the most important functions of the science of paleontology. Such a preserved specimen is called a “fossil” if it is older than some minimum age, most often the arbitrary date of 10,000 years ago. Hence, fossils range in age from the youngest at the start of the Holocene Epoch to the oldest from the Archaean Eon, up to 3.4 billion years old. The observations that certain fossils were associated with certain rock strata led early geologists to recognize a geological timescale in the 19th century. The development of radiometric dating techniques in the early 20th century allowed geologists to determine the numerical or “absolute” age of the various strata and thereby the included fossils.

Like extant organisms, fossils vary in size from microscopic, such as single bacterial cells[4] only one micrometer in diameter, to gigantic, such as 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. Preservation of soft tissues is rare in the fossil record. Fossils may also consist of the marks left behind by the organism while it was alive, such as the footprint or feces (coprolites) of a reptile. These types of fossil are called trace fossils (or ichnofossils), as opposed to body fossils. Finally, past life leaves some markers that cannot be seen but can be detected in the form of biochemical signals; these are known as chemofossils or biomarkers.

Interpretation of the fossil record

Ever since recorded history began, and probably before, people have noticed and gathered fossils, including pieces of rock and minerals that have replaced the remains of biologic organisms, or preserved their external form. Fossils themselves, and the totality of their occurrence within the sequence of Earth’s rock strata, is referred to as the fossil record.

The fossil record was one of the early sources of data relevant to the study of evolution and continues to be relevant to the history of life on Earth. Paleontologists examine the fossil record in order to understand the process of evolution and the way particular species have evolved.

Various explanations have been put forth throughout history to explain what fossils are and how they came to be where they were found. Many of these explanations relied on folktales or mythologies. In China the fossil bones of ancient mammals including Homo erectus were often mistaken for “dragon bones” and used as medicine and aphrodisiacs. In the West the presence of fossilized sea creatures high up on mountainsides was seen as proof of the biblical deluge.

Greek scholar Aristotle realized that fossil seashells from rocks were similar to those found on the beach, indicating the fossils were once living animals.Leonardo da Vinci concurred with Aristotle’s view that fossils were the remains of ancient life. In 1027, the Persian geologist, Avicenna explained how thestoniness of fossils was caused in The Book of Healing. However, he rejected the explanation of fossils as organic remains. Aristotle previously explained it in terms of vaporous exhalations, which Avicenna modified into the theory of petrifying fluids (succus lapidificatus), which was elaborated on by Albert of Saxony in the 14th century and accepted in some form by most naturalists by the 16th century. Avicenna gave the following explanation for the origin of fossils from thepetrifaction of plants and animals:

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.

More scientific views of fossils emerged during the Renaissance. For example, Leonardo Da Vinci noticed discrepancies with the use of 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.”

Biological explanations

Early naturalists well understood the similarities and differences of living species leading Linnaeus to develop a hierarchical classification system still in use today. It was Darwin and his contemporaries who first linked the hierarchical structure of the great tree of life in living organisms 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 Charles 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. The absence of older fossils worried Darwin about the implications for 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.

Rare Fossils

Fossilization is an exceptionally rare occurrence, because most components of formerly living things tend to decompose relatively quickly following death. In order for an organism to be fossilized, the remains normally need to be covered by sediment as soon as possible. However there are exceptions to this, such as if an organism becomes frozen, desiccated, or comes to rest in an anoxic (oxygen-free) environment. There are several different types of fossils and fossilization processes.

Due to the combined effect of taphonomic processes and simple mathematical chance, fossilization tends to favor organisms with hard body parts, those that were widespread, and those that existed for a long time before going extinct. On the other hand, it is very unusual to find fossils of small, soft bodied, geographically restricted and geologically ephemeral organisms, because of their relative rarity and low likelihood of preservation.

Larger specimens (macrofossils) are more often observed, dug up and displayed, although microscopic remains (microfossils) are actually far more common in the fossil record.

Some casual observers have been perplexed by the rarity of transitional species within the fossil record. The conventional explanation for this rarity was given by Darwin, who stated that “the extreme imperfection of the geological record,” combined with the short duration and narrow geographical range of transitional species, made it unlikely that many such fossils would be found. Simply put, the conditions under which fossilization takes place are quite rare; and it is highly unlikely that any given organism will leave behind a fossil. Eldredge and Gould developed their theory of punctuated equilibrium in part to explain the pattern of stasis and sudden appearance in the fossil record. Furthermore, in the strictest sense, nearly all fossils are “transitional,” due to the improbability that any given fossil represents the absolute termination of an evolutionary path.

Types of Preservation

1. Permineralization

Permineralization occurs after burial, as the empty spaces within an organism (spaces filled with liquid or gas during life) become filled with mineral-rich groundwater and the minerals precipitate from the groundwater, thus 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 decaying 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.

2. Casts and molds

In some cases the original remains of the organism have been completely dissolved or otherwise destroyed. When all that is left is an organism-shaped hole in the rock, it 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.

3. Authigenic mineralisation

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 authigenic mineralisation.

4.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.

5. Adpression (compression-impression) fossils

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, it has proved to convenient to have a combined term for both modes of preservation: adpression.

6. Bioimmuration

Bioimmuration is a type of preservation in which a skeletal organism overgrows or otherwise subsumes another organism, preserving the latter, or an impression of it, within the skeleton.] Usually it is a sessile skeletal organism, such as a bryozoan or an oyster, which grows along a substrate, covering other sessile encrusters. 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 and grows upwards, preserving the settler in its skeleton. Bioimmuration is known in the fossil record from the Ordovicianto the Recent. To sum up, fossilization processes proceed differently for different kinds of tissues and under different kinds of conditions.

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