Concepts of taphonomy and biostratinomy

TAPHONOMY
Taphonomy is the study of what happens to an organism after its death and until its discovery as a fossil. This includes decomposition, post-mortem 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 life-history of the once-living 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 post-mortem 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 shell-hash or coquina  has experienced a significant amount of shell breakage and probably post-mortem transport suggesting deposition in high energy environments; whereas, the articulated plant remains   are intact suggesting little or no post-mortem transport and deposition in a very low energy and oxygen-free 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 TheirPaleoenvironmental Implications
TAPHONOMIC FEATURE
IMPLICATIONS
Abrasion The wearing-down of skeletons owing to differential movement with respect to sediments is an indicator of environmental energy. Significant abrasion is most commonly found on skeletal material collected from beaches, or areas of strong currents or wave action.
Articulation Multi-element skeletons are soon disarticulated after death. Articulated skeletons, then, indicate rapid burial or otherwise removing the skeleton from the effects of energy of the original environment.
Bioerosion 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 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.
Dissolution Skeletal remains commonly are in equilibrium with surrounding waters, but changes in chemical conditions can cause skeletons to dissolve. Dissolution represents fluctuation 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.
Rounding Broken edges of skeletons become rounded owing to dissolution and/or abrasion of exposed surfaces. Processes that control edge rounding probably include a combination of dissolution, abrasion, and bioerosion. Rounding gives an estimate of time since breakage.
Encrustation The growth of hard skeleton substrates by other organisms is a common occurrence. Besides indicating exposure of the skeleton above the sediment-water interface, encrustation can specify a particular environment. Different patterns of encrustation, as well as different biota, occur in different environments.
Fragmentation Breakage of skeletons is usually an indication of high energy resulting from wave action or current energy. 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 in-place infauna. Hard parts tend to orient long-axis parallel to unidirectional flow in current-dominated areas and perpendicular to wave crests on wave-dominated bottoms.
Size After death and if not rapidly buried, a skeleton behaves as a sedimentary particle and is moved and sorted with respect to 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 wind-driven system.

From McRoberts (1998)

 



A Brief Introduction to

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.


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