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TAPHONOMY
©
Gastaldo, Savrda, & Lewis. 1996. Deciphering Earth History: A
Laboratory Manual with Internet Exercises. Contemporary
Publishing Company of Raleigh, Inc. ISBN 0-89892-139-2
Not every organism that ever lived could become part of the fossil record. If you eat an
average of three meals a day, you test and prove this hypothesis daily. A large
percentage of all biological entities end up as food for other organisms higher on the
food chain. This fact alone may prevents these organisms from being preserved. Even
those organisms that avoid being eaten have a low probability of becoming fossilized
because most of them undergo decay and recycling of their chemical components. For
example, you can examine any forest-floor litter and find that beneath the top layer of
leaves, the organic matter has been degraded to an unrecognizable form (humus -- not
hummus, the garlic-laden spread served in health-food restaurants). This recycling
keeps the carbon, nitrogen, and sulfur cycles operating. In fact, many taphonomic biases
impact the odds of any organism being preserved.
The paleontological subdiscipline called Taphonomy, from the Greek taphos (death), is
concerned with the processes responsible for any organism becoming part of the fossil
record and how these processes influence information in the fossil record. Many
taphonomic processes must be considered when trying to understand fossilization.
These include events that affected the organism during life (changes in rainfall,
availability of food, and behavior for maximum growth, etc.), the transferral of that
organism (or a part of that organism) from the living world (biosphere) to the
sedimentary record (lithosphere; compare the death of a herd of vertebrates with the
autumnal leaf fall from a forest), and the physical and chemical interactions that affect
the organism from the time it is buried until the time it is collected in the field.
Any organism must successfully pass through three distinct, and separate, stages in
order to be seen in a museum display. These stages span the entire time from death of
the organism to collection. Necrology is the first stage, and involves the death or loss of
a part of the organism. The vast majority of animals must die before they can become
introduced to the next phase. It's true that if a starfish is cut in half, each half will
regenerate itself. The result will be two animals. Not many animals have this capability.
We suggest that you don't test this hypothesis with your beloved pet. On the other hand,
most plants do not have to die to contribute one or more of their parts to the potential
fossil record. When autumn leaves fall in temperate climates, the trees don't die. The
oldest living organism, bristlecone pines, are more than 5300 years old (as determined
by counting tree rings). Their present leaves are not the same ones that grew 5300 years
ago. When plants disperse their reproductive bodies (spores, pollen, or seeds), most do
not die thereafter. Of course there are exceptions, but these are a small percentage of all
extant (living) plants.
Once an organism has died or sheds a part, all the interactions involving its transferral
from the living world to the inorganic world (including burial) constitute the second
taphonomic stage. This is the Biostratinomy stage. Besides the conspicuous fossil
characteristics that you will be able to observe during this laboratory (those external and
internal features of the fossilized remain), less-obvious details often record what
happened to the organism (or part) before it became a fossil. By studying these details
paleontologists are able to understand, in a Sherlock Holmesian way, the mode of death
or disarticulation (breakup of an organism), any biological processes that may have
modified the remains before burial (such as scavenging), the response of the part to
transport (by animals, water and/or wind), and the amount of time the organism sat
around in the environment before it was finally entombed.
Ultimately the organic matter is buried. Burial plays an important role in potential
preservation of the organic matter. Very specific chemical and physical conditions must
exist in the burial environment to allow preservation in a form recognizable to us. It is
here that biological (e.g., enzymatic and bacterial) and chemical (e.g., enzymatic and
dissolution) processes must be slowed or eliminated. Once buried, the organic material
is subjected to the third taphonomic phase, or Diagenesis. Diagenesis involves all of the
processes responsible for lithification of the sediment and chemical interactions with
waters residing between clasts. The processes of fossilization appear to be site specific
with respect to depositional settings, resulting in a mosaic of preservational traits in the
terrestrial and marine realm. Few fossil assemblages are exactly identical, especially
with regard to the way in which they were formed, but general patterns do exist. An
understanding of taphonomic assemblage features within an environmental context
allows for a more accurate interpretation of the fossil record.
Most organic matter on Earth is used by some organism higher on the food chain and is,
therefore, ultimately recycled. This is the fate of almost all biomass on Earth. Most
organic matter is composed of easily degraded and digested compounds that are not
likely to be preserved even under the most favorable conditions. Those parts of an
organism that are already mineralized (such as your calcium-fortified skeleton) and,
hence have made the first step in the transition to "stone", have a higher probability of
preservation than any of the soft, fleshy tissues either around or within the skeleton. The
early inhabitants of Paris, France, the bones of whom are now stacked neatly in
catacombs beneath the city streets, attest to this fact.
Although the fossil record is incomplete, it still provides a useful survey of the history
of life because of the vast amounts of time represented within the rock record. Even if
the conditions for preserving organic matter existed only once every 10,000 years in
each contemporaneous depositional environment around the globe, a lithology that was
100 meters thick (330 feet) and encompassing 1 million years of time would contain
100 fossil assemblages. Such conditions are not unrealistic, particularly within the
ocean basins. If we then consider contemporaneous depositional settings around the
globe, the number of fossil assemblages that would be preserved during this 1 million
years of time increases dramatically. Of course, not all of these fossil sites are or would
be accessible for collection and study. Mountain-building processes associated with
plate tectonic activity (metamorphism of fossil-bearing sedimentary rock beyond
recognition) and the erosion of these folded (metasedimentary) and faulted
(sedimentary) rocks depletes the number of fossil localities available at the Earth's
surface through time. The quantity of fossiliferous rocks beneath ground level far
exceeds those available at the surface to be sampled and studied. Nevertheless, there are
far more fossils than paleontologists, which will continue to be the case far into the
future. Paleontologists are not wanting in their search for the history of life on Earth.
The next 40 years was a period of expanding research on the nature and behavior of
atoms, leading to the development of nuclear fission and fusion as energy sources. A
byproduct of this atomic research has been the development and continuing refinement
of the various methods and techniques used to measure the age of Earth materials.
Precise dating has been accomplished since 1950.
A chemical element consists of atoms with a specific number of protons in their nuclei
but different atomic weights owing to variations in the number of neutrons. Atoms of
the same element with differing atomic weights are called isotopes. Radioactive decay
is a spontaneous process in which an isotope (the parent) loses particles from its nucleus
to form an isotope of a new element (the daughter). The rate of decay is conveniently
expressed in terms of an isotope's half-life, or the time it takes for one-half of a
particular radioactive isotope in a sample to decay. Most radioactive isotopes have rapid
rates of decay (that is, short half-lives) and lose their radioactivity within a few days or
years. Some isotopes, however, decay slowly, and several of these are used as geologic
clocks. The parent isotopes and corresponding daughter products most commonly used
to determine the ages of ancient rocks are listed below:
The mathematical expression that relates radioactive decay to geologic time is called the
age equation and is:
Dating rocks by these radioactive timekeepers is simple in theory, but the laboratory
procedures are complex. The numbers of parent and daughter isotopes in each specimen
are determined by various kinds of analytical methods. The principal difficulty lies in
measuring precisely very small amounts of isotopes.
The potassium-argon method can be used on rocks as young as a few thousand years as
well as on the oldest rocks known. Potassium is found in most rock-forming minerals,
the half-life of its radioactive isotope potassium-40 is such that measurable quantities of
argon (daughter) have accumulated in potassium-bearing minerals of nearly all ages,
and the amounts of potassium and argon isotopes can be measured accurately, even in
very small quantities. Where feasible, two or more methods of analysis are used on the
same specimen of rock to confirm the results.
Another important atomic clock used for dating purposes is based on the radioactive
decay of the isotope carbon-14, which has a half-life of 5,730 years. Carbon-14 is
produced continuously in the Earth's upper atmosphere as a result of the bombardment
of nitrogen by neutrons from cosmic rays. This newly formed radiocarbon becomes
uniformly mixed with the nonradioactive carbon in the carbon dioxide of the air, and it
eventually finds its way into all living plants and animals. In effect, all carbon in living
organisms contains a constant proportion of radiocarbon to nonradioactive carbon. After
the death of the organism, the amount of radiocarbon gradually decreases as it reverts to
nitrogen-14 by radioactive decay. By measuring the amount of radioactivity remaining
in organic materials, the amount of carbon-14 in the materials can be calculated and the
time of death can be determined. For example, if carbon from a sample of wood is
found to contain only half as much carbon-14 as that from a living plant, the estimated
age of the old wood would be 5,730 years.
The radiocarbon clock has become an extremely useful and efficient tool in dating the
important episodes in the recent prehistory and history of man, but because of the
relatively short half-life of carbon-14, the clock can be used for dating events that have
taken place only within the past 50,000 years.
The following is a group of rocks and materials that have dated by various atomic clock
methods:
Approximate Age
Sample
in Years
Cloth wrappings from a mummified bull
Samples taken from a pyramid in Dashur, Egypt. This date
2,050
agrees with the age of the pyramid as estimated from historical
records
Charcoal
Sample, recovered from bed of ash near Crater Lake, Oregon,
is from a tree burned in the violent eruption of Mount Mazama
6,640
which created Crater Lake. This eruption blanketed several
States with ash, providing geologists with an excellent time
zone.
Charcoal
Sample collected from the "Marmes Man" site in southeastern
10,130
Washington. This rock shelter is believed to be among the
oldest known inhabited sites in North America
Spruce wood
Sample from the Two Creeks forest bed near Milwaukee,
11,640
Wisconsin, dates one of the last advances of the continental ice
sheet into the United States.
Bishop Tuff
Samples collected from volcanic ash and pumice that overlie
glacial debris in Owens Valley, California. This volcanic 700,000
episode provides an important reference datum in the glacial
history of North America.
Volcanic ash
Samples collected from strata in Olduvai Gorge, East Africa,
1,750,000
which sandwich the fossil remains of Zinjanthropus and Homo
habilis -- possible precursors of modern man.
Monzonite
Samples of copper-bearing rock from vast open-pit mine at 37,500,000
Bingham Canyon. Utah.
Quartz monzonite
Samples collected from Half Dome, Yosemite National Park, 80,000,000
California.
Conway Granite
Samples collected from Redstone Quarry in the White 180,000,000
Mountains of New Hampshire.
Rhyolite
Samples collected from Mount Rogers, the highest point in 820,000,000
Virginia.
Pikes Peak Granite
1,030,000,000
Samples collected on top of Pikes Peak, Colorado.
Gneiss
Samples from outcrops in the Karelian area of eastern Finland 2,700,000,000
are believed to represent the oldest rocks in the Baltic region.
The Old Granite
Samples from outcrops in the Transvaal, South Africa. These 3,200,000,000
rocks intrude even older rocks that have not been dated.
Morton Gneiss [see Editor's Note] 3,600,000,000
Samples from outcrops in southwestern Minnesota are
believed to represent some of the oldest rocks in North
America.
Interweaving the relative time scale with the atomic time scale poses certain problems
because only certain types of rocks, chiefly the igneous variety, can be dated directly by
radiometric methods; but these rocks do not ordinarily contain fossils. Igneous rocks are
those such as granite and basalt which crystallize from molten material called "magma".
When igneous rocks crystallize, the newly formed minerals contain various amounts of
chemical elements, some of which have radioactive isotopes. These isotopes decay
within the rocks according to their half-life rates, and by selecting the appropriate
minerals (those that contain potassium, for instance) and measuring the relative amounts
of parent and daughter isotopes in them, the date at which the rock crystallized can be
determined. Most of the large igneous rock masses of the world have been dated in this
manner.
Most sedimentary rocks such as sandstone, limestone, and shale are related to the
radiometric time scale by bracketing them within time zones that are determined by
dating appropriately selected igneous rocks, as shown by a hypothetical example.
Literally thousands of dated materials are now available for use to bracket the various
episodes in the history of the Earth within specific time zones. Many points on the time
scale are being revised, however, as the behavior of isotopes in the Earth's crust is more
clearly understood. Thus the graphic illustration of the geologic time scale, showing
both relative time and radiometric time, represents only the present state of knowledge.
Certainly, revisions and modifications will be forthcoming as research continues to
improve our knowledge of Earth history.
http://www.ucmp.berkeley.edu/IB181/VPL/Pres/Pres4.html
The study of how organisms or their parts become fossils is called taphonomy.
Taphonomy is literally everything that happens to an organism - or part of an
organism, as is often the case with plants - from the moment that it dies until it is
collected and curated for scientific study. Figure 3.1 illustrates some of the many
taphonomic pathways a plant or plant part can take from it's living community to
the museum drawer.
Levels of preservation
Plant fossil preservation can take place at a number of levels. Each level contains a
different type of information.
Cellular Level
Not all organic compounds are equally resistant to chemical degradation and decay.
Plant cell walls (composed primarily of the polysaccharide polymer cellulose) are far
more likely to escape decomposition than internal membranes and organelles,
which are rich in protein, lipids and sugars. Preservation of cytological details has
been reported in fossil plants, but occurrences are rare, and most reports of
fossilized nuclei and organelles should be read with caution. Secondary compounds,
such as those impregnating or covering cell walls, can also be resistant to
decomposition; examples include lignin, waxes, cutin (which comprises plant
cuticle), and sporopollenin, which forms the external shell of spores, pollen, and the
resting cysts of some marine algae.
Tissue Level
Organ Level
Plants break apart, both in life (organ senescence or dispersal) and after death;
dispersed parts may be transported before settling into the sediment to be buried
and become fossils. Assemblages of plant fossils that are preserved close to where
their parent-plants originally grew are called autochthonous; assemblages that
have been transported are referred to as allochthonous. Whether an assemblage is
autochthonous or transported has obvious implications for what sorts of ecological
interpretations we can make from it. Therefore, as you study each type of plant
fossil preservation, think about whether fossils produced in these ways are
autochthonous or not.
Because we don't always know which leaves belong to which seeds when they are
first discovered, we use the convention of form taxa. When organs are found
isolated (not in organic connection), each type of leaf and seed is given it's own
binomial name (genus and species name according to the International Code of
Botanical Nomenclature), without making any assumption about what belongs to
what. To use the example discussed by Oliver and Scott (1904), leaves were
described as Lyginopteris (genus only for brevity), seeds as Lagenostoma, and
stems as Lyginodendron. The similarity of the first syllable gives a hint that the
describing paleobotanists (others besides Oliver and Scott) suspected some
relationship, but were unable to make a strong inference link. The last syllable of
each name gives a hint to the organ type: "dendron" = stem, "pteris" is often used
for frond-like foliage, "stoma" = seed. However, after Oliver and Scott's recognition
of the unique glands on Lagenostoma lomaxi and species in the other organ form
genera, they were able to make the whole-plant link with greater confidence. The
whole plant then takes the name of the organ first described, in this
caseLyginopteris. When you are writing, take care to make clear whether you are
talking about form taxa (organs) or whole-plants.
Form taxa are not unique to paleobotany. For example, the planktonic larvae of
many marine invertebrates are commonly described as separate species when they
are first discovered in the ocean. Only later when they can be reared in the
laboratory can the link to their adult form be recognized. Similarly, the different life
stages of many fungi are given different names because they have different
physical forms and hosts. Only through detailed inoculation studies can mycologists
work out which forms are members of the same life cycle. Since some fungi may
have more than five discrete life cycle stages, this can be a long process. Similar
problems exist for some marine algae and multiple-host parasitic organisms of
many kinds. Even among well-studied vertebrates, some tropical birds have been
described as separate species until they are observed to mate and rear young
together.
Organism Level
Not all plants in a given community are equally likely to find their way into the fossil
record. Processes of fossilization often favor large or woody plants with resistant
tissues over small herbs. Likewise, wind dispersed pollen is much more common in
the fossil record than pollen dispersed by animals. Also, plants growing in or near
an area of preservation (e.g., riverbank or swamp) are more commonly preserved
than their counterparts growing far from water or anoxic sedimentary
environments.
Environmental Level
Plant preservation depends on removing the organic material from the zone of
aerobic decomposition. This is most easily accomplished by burying the plant.
Consequently, swamps, deltas, lakes, lowland flood plains, and volcanic areas are
good spots for fossilization (VG 1:9)(VG 1:12)(VG 1:13). Arid regions and
mountainsides are not likely candidates for plant fossil preservation (with the
exception of exquisitely preserved plant material from Pleistocene packrat middens
in the southwestern United States.
All of these taphonomic factors influence the information that can be recovered
from the plant fossil record. While taphonomic filtering does not preclude biological
interpretation of fossil material, taphonomy can introduce substantial biases into
the record and influence our interpretation of the fossils, and thus our
reconstruction of the ancient plants. It is, therefore, important always to keep in
mind the mode of preservation of fossils when making any interpretations from
fossils.
Impressions
Permineralizations
Compactions
Molecular Fossils
Fossil DNA and RNA have also been making headlines in the scientific
press. In some exceptional cases, genetic material or proteins have
been sufficiently well-preserved to permit their use in the
reconstruction of evolutionary relationships, in much the same way as
one might sequence living organisms. However, much of this work is
controversial due to the difficulty of preserving and isolating these
fragile molecules. Also, contamination by other materials is a
common and difficult to recognize problem.
Moldowan, J.M., J. Dahl, B.J. Huizinga, F.J. Fago, L.J. Hickey, T.M.
Peakman, and D.W. Taylor. 1994. The molecular fossil record of
loeanane and its relation to angiosperms. Science 265:768-771.
Oliver, F.W. and D.H. Scott. 1904. On the structure of the Palaeozoic
seed Lagenostoma lomaxi, wit a statment of the evidence upon which
it is referred to Lyginodendron. Philosophical Transactions of the
Royal Society of London 197B: 193-248.
TAPHONOMY & PRESERVATION
INTRODUCTION
As fossils are the preserved remains of ancient organisms or their traces,
understanding the process of preservation, and more importantly, being able to
recognize and identify fossil remains after their discovery is an integral part of
paleobiology. Protective cover (sediments) and stabilizing chemical environments
are of prime importance in the preservation of once living organisms. Due to the
process of aerobic decay and physical/chemical destruction, most animals leave no
evidence of their existence.
In order to make a correct interpretation of taphonomic processes and mode of
preservation, it is often necessary to have a prior knowledge of the structural
features or morphology of original skeleton in addition to knowing its original
mineralogical composition. This limitation should diminish as you become
familiar with the various fossil groups throughout the semester.
TAPHONOMY
Taphonomy is the study of what happens to an organism after its death and until
its discovery as a fossil. This includes decomposition, postmortem transport,
burial, compaction, and other chemical, biologic, or physical activity which affects
the remains of the organism. Being able to recognize taphonomic processes that
have taken place can often lead to a better understanding of paleoenvironments
and even lifehistory of the onceliving organism.
In addition, understanding which taphonomic processes a fossil occurrence has
undergone, and to what degree, may have implication on interpreting the
significance of the fossil deposit and clearer understanding of the biases in the
sample.
An outline of the pathways affecting the preservation of once living organisms can
be found in Figure 1 below. As discussed below, this encompasses both the
processes of biostratinomy and diagenesis.
Figure 1 - The field of Taphonomy as it relates to steps in
transformation from living organisms to fossils.
Modified from McRoberts (1998)
Processes that occur between the death of an organism and its subsequent burial in
the sediment are termed biostratinomy. Generally, this includes the
decomposition and scavenging of the animal's soft parts, and at least some amount
of postmortem transport. Such things as the amount of shell breakage and the
concentration of shells in layers often indicate the level of water energy and post
mortem transport. For example, the shellhash or coquina has experienced a
significant amount of shell breakage and probably postmortem transport
suggesting deposition in high energy environments; whereas, the articulated plant
remains are intact suggesting little or no postmortem transport and deposition
in a very low energy and oxygenfree environment. In Table 1 below are various
taphonomic indicators and their environmental implications.
The physical and/or chemical effects after burial are called diagenesis. This is the
realm in which dissolution, replacement, or recrystallization of original shell
material occurs, as can the formation of molds and casts. A more detailed
description of diagenesis with regards to fossil preservation in the next section.
Table 1
Summary of Taphonomic Indicators and Their Paleoenvironmental
Implications
TAPHONOMIC
IMPLICATIONS
FEATURE
The wearingdown of skeletons owing to differential
movement with respect to sediments is an indicator of
Abrasion environmental energy. Significant abrasion is most commonly
found on skeletal material collected from beaches, or areas of
strong currents or wave action.
Multielement skeletons are soon disarticulated after death.
Articulated skeletons, then, indicate rapid burial or otherwise
Articulation
removing the skeleton from the effects of energy of the
original environment.
Bioerosion encompasses the many different corrosive
processes by organisms. The most pervasive causes of
degradation are boring and grazing. Bioerosion erases
information from the fossil record, but it also leaves
Bioerosion
identifiable traces made by organisms on remaining hard
skeletons or surfaces. Therefore, trace fossils produced by
bioerosion add information on the diversity of ancient
assemblages.
Skeletal remains commonly are in equilibrium with
surrounding waters, but changes in chemical conditions can
cause skeletons to dissolve. Dissolution represents fluctuation
Dissolution
in temperature, pH or pCO2 in calcium carbonate skeletons.
Siliceous skeletons also can dissolve because normal sea water
is usually undersaturated with respect to silica.
Broken edges of skeletons become rounded owing to
dissolution and/or abrasion of exposed surfaces. Processes
Rounding that control edge rounding probably include a combination of
dissolution, abrasion, and bioerosion. Rounding gives an
estimate of time since breakage.
The growth of hard skeleton substrates by other organisms is a
common occurrence. Besides indicating exposure of the
skeleton above the sedimentwater interface, encrustation can
Encrustation
specify a particular environment. Different patterns of
encrustation, as well as different biota, occur in different
environments.
Breakage of skeletons is usually an indication of high energy
resulting from wave action or current energy. Fragmentation
Fragmentation
also can be caused by other organisms through either
predation or scavenging.
Orientation After death, skeletal remains are moved by the transporting
medium and oriented relative to their hydrodynamic
properties. Fossil skeletons in life position indicate rapid
burial, attachment to a firm substrate, or death of inplace
infauna. Hard parts tend to orient longaxis parallel to
unidirectional flow in currentdominated areas and
perpendicular to wave crests on wavedominated bottoms.
After death and if not rapidly buried, a skeleton behaves as a
sedimentary particle and is moved and sorted with respect to
Size the carrying capacity of the flow of currents, waves, or tides.
Size can, therefore, be an effective indicator of flow capacity in
a hydraulic or winddriven system.
From McRoberts (1998)
FORMS OF PRESERVATION
UNALTERED
This form of preservation is rare in most of the geologic column, but becomes more
frequent in younger sedimentary rocks. Types of unaltered preservation where
even the soft body parts are preserved include: (i) mummification, (ii) encasement
in tar, (iii) encasement in amber, (iv) encasement in sediment, and (v) freezing.
More frequently, however, only the hard skeletal material is preserved after
removal of soft body parts.
Examples of unaltered preservation include the skeleton of a horseshoe crab,
whose shell is composed of interlocking plates and jointed appendages which
quickly disarticulate after death; cockle bivalved molluscs, whose outermost
shell layer has been removed by abrasion, yet the original shell material of the
inner layers remains; an ammonoid from the Cretaceous period in which you
should note the pearly luster which is original aragonite shell material; and an
insect encased in amber .
MOLDS & CASTS
This general class of preservation entails making "replicas" of the skeletal hard
parts of organisms. In general, a mold is an impression in the sediment of a
skeleton or shell. Once encased in lithified sediment, the dissolution of skeletal
material leaves behind the impression or mold of original skeletal form. Thus, a
mold is a "mirror image" of the original skeleton. An internal mold (sometimes
called a steinkern) is the impression of the inside surface of skeletal hard parts. An
external mold is the impression of the outside surface of skeleton or bone. An
example of both types of molds can be seen in this image of a trilobite .
A cast is formed by the fillingin of a mold. It is thus a true replica (not a "mirror
image") of the original skeleton or shell. By this definition, the cast one gets for a
broken limb is not really a cast at all but an external mold.
A graphical representation of the formation of casts and molds is provided in
Figure 2 below.
Figure 2 Different diagenetic processes leading to different
preservational styles in skeletal materials.
* Note that molds are produced directly as imprints of the shell and casts are produced from
molds.
REPLACEMENT & RECRYSTALLIZATION
This common form of preservation involves chemical and/or physical alteration or
replacement of original skeletal material. To properly identify replacement and
recrystallization, one must know what the original constituents of the organism's
skeleton were. These are provided in Figure 1.3. Replacement occurs often by the
filling in (by various minerals) of the void space after dissolution of original
skeletal material. Sometimes, the replacement occurs on a molecule by molecule
basis. Common replacement minerals that you should be able to recognize include
Silica (SiO2) as shown in the coral, and Pyrite (FeS2) shown in the ammonoid.
Recrystallization involves the physical rearrangement of crystalline structure of
skeletal material. This is a common phenomenon in shells which were originally
aragonite and/or calcite (both forms of calcium carbonate CaCO3). Examples,
both of which are now calcite, include a gastropod which was originally aragonite
and a brachiopod which was originally calcite.
CARBONIZATION
As organic remains decompose in sediments, volatile constituents such as oxygen,
hydrogen, and nitrogen are slowly lost to the surrounding sediments frequently
leaving behind a carbon film. This process is carbonization (or sometimes called
distillation), and occurs most frequently in oxygen deficient, organicrich
environments such as basinal black shales, and coal swamps. The carbon films
often show exquisite details of plants and softbody parts of animals not readily
preserved, and can often be recognized by a dark gray or black film with a metallic
sheen such as these fernlike fossil plants.
PERMINERALIZATION
Permineralization involves the fillingin of pore and/or void spaces in shell or
bone by secondary mineral matter in solution. With permineralization, the tiny
pore spaces in the fossil are filled and the original skeletal material is still retained.
However, it is often common for other types of preservation (e.g. replacement) to
occur during and/or after permineralization. Because of its porous nature, bone
and wood is especially prone to permineralization.