Taphonomy is the study of how organisms decay and become fossilized.
The term taphonomy (from the Greek taphos, τάφος meaning
"burial", and nomos, νόμος meaning "law") was introduced to
paleontology in 1949 by Russian scientist Ivan Efremov to describe
the study of the transition of remains, parts, or products of
organisms from the biosphere to the lithosphere.
2 Research areas
2.3 Forensic science
2.4 Environmental archaeology
3 Taphonomic biases in the fossil record
3.1 Physical attributes of the organism itself
3.2 Characteristics of the habitat
3.3 Mixing of fossils from different places
3.4 Temporal resolution
3.5 Gaps in time series
3.6 Consistency in preservation over geologic time
3.7 Human biases
4 Preservation of biopolymers
7 See also
9 Further reading
10 External links
Taphonomic phenomena are grouped into two phases: biostratinomy;
events that occur between death of the organism and the burial, and
diagenesis; events that occur after the burial. Since Efremov's
definition, taphonomy has expanded to include the fossilization of
organic materials, inorganic materials, and both cultural and
This is a multidisciplinary concept and is used in slightly different
contexts throughout different fields of study. Fields that employ the
concept of taphonomy include:
An articulated wombat skeleton in Imperial-Diamond cave (Jenolan
La Brea Tar Pits
La Brea Tar Pits represent an unusual depositional environment for
their epoch (Pleistocene) and location (southern California).
There are five main stages of taphonomy: disarticulation, dispersal,
accumulation, fossilization, and mechanical alteration. The first
stage, disarticulation, occurs as the organism decays and the bones
are no longer held together by the flesh and tendons of the organism.
Dispersal is the separation of pieces of an organism caused by natural
events (i.e. floods, scavengers etc.). Accumulation occurs when there
is a buildup of organic and/or inorganic materials in one location
(scavengers or human behavior). When mineral rich groundwater
permeates organic materials and fills the empty spaces, a fossil is
formed. The final stage of taphonomy is mechanical alteration; this is
processes that physically alter the remains (i.e. freeze-thaw,
compaction, transport, burial).
Taphonomy has undergone an explosion of interest since the 1980s,
with research focusing on certain areas.
Microbial, biogeochemical, and larger-scale controls on the
preservation of different tissue types; in particular, exceptional
preservation in Konzervat-lagerstätten. Covered within this field is
the dominance of biological versus physical agents in the destruction
of remains from all major taxonomic groups (plants, invertebrates,
Processes that concentrate biological remains; especially the degree
to which different types of assemblages reflect the species
composition and abundance of source faunas and floras.
The spatio-temporal resolution[clarification needed] and ecological
fidelity[clarification needed] of species assemblages, particularly
the relatively minor role of out-of-habitat transport contrasted with
the major effects of time-averaging.[clarification needed]
The outlines of megabiases in the fossil record, including the
evolution of new bauplans and behavioral capabilities, and by
broad-scale changes in climate, tectonics, and geochemistry of Earth
Mars Science Laboratory
Mars Science Laboratory mission objectives evolved from assessment
Mars habitability to developing predictive models on
One motivation behind taphonomy is to better understand biases present
in the fossil record.
Fossils are ubiquitous in sedimentary rocks, yet
paleontologists cannot draw the most accurate conclusions about the
lives and ecology of the fossilized organisms without knowing about
the processes involved in their fossilization. For example, if a
fossil assemblage contains more of one type of fossil than another,
one can infer either that the organism was present in greater numbers,
or that its remains were more resistant to decomposition.
During the late twentieth century, taphonomic data began to be applied
to other paleontological subfields such as paleobiology,
paleoceanography, ichnology (the study of trace fossils) and
biostratigraphy. By coming to understand the oceanographic and
ethological implications of observed taphonomic patterns,
paleontologists have been able to provide new and meaningful
interpretations and correlations that would have otherwise remained
obscure in the fossil record.
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
On January 24, 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. The search for evidence of habitability,
taphonomy (related to fossils), and organic carbon on the planet Mars
is now a primary
Forensic taphonomy is a relatively new field that has increased in
popularity in the past 15 years. It is a subfield of forensic
anthropology focusing specifically on how taphonomic forces have
altered criminal evidence.
There are two different branches of forensic taphonomy: biotaphonomy
and geotaphonomy. Biotaphonomy looks at how the decomposition and/or
destruction of the organism has happened. The main factors that affect
this branch are categorized into three groups: environmental factors;
external variables, individual factors; factors from the organism
itself (i.e. body size, age, etc.), and cultural factors; factors
specific to any cultural behaviors that would affect the decomposition
(burial practices). Geotaphonomy studies how the burial practices and
the burial itself affects the surrounding environment. This includes
soil disturbances and tool marks from digging the grave, disruption of
plant growth and soil pH from the decomposing body, and the alteration
of the land and water drainage from introducing an unnatural mass to
This field is extremely important because it helps scientists use the
taphonomic profile to help determine what happened to the remains at
the time of death (perimortem) and after death (postmortem). This can
make a huge difference when considering what can be used as evidence
in a criminal investigation.
Archaeologists study taphonomic processes in order to determine how
plant and animal (including human) remains accumulate and
differentially preserve within archaeological sites. Environmental
Archaeology is a multidisciplinary field of study that focuses on
understanding the past relationships between groups and their
environments. The main subfields of environmental archaeology include
zooarchaeology, paleobotany, and geoarchaeology.
specialists to identify what artifacts or remains encountered before
and after initial burial. Zooarchaeology, a focus within environmental
archaeology investigates taphonomic processes on animal remains. The
processes most commonly identified within zooarchaeology include
thermal alteration (burns), cut marks, worked bone, and gnaw
marks. Thermally altered bone indicate the use of fire and animal
processing. Cut marks and worked bone can inform zooarchaeologists on
tool use or food processing. When there is little to no written
record, taphonomy allows environmental archaeologists to better
comprehend the ways in which a group interacted with their surrounding
environments and inhabitants.
The field of environmental archaeology provides crucial information
for attempting to understand the resilience of past societies and the
great impacts that environmental shifts can have on a population.
Knowledge gained from the past through these studies can be used to
inform present and future decisions for human-environment
Taphonomic biases in the fossil record
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Because of the very select processes that cause preservation, not all
organisms have the same chance of being preserved. Any factor that
affects the likelihood that an organism is preserved as a fossil is a
potential source of bias. It is thus arguably the most important goal
of taphonomy to identify the scope of such biases such that they can
be quantified to allow correct interpretations of the relative
abundances of organisms that make up a fossil biota. Some of the
most common sources of bias are listed below.
Physical attributes of the organism itself
This perhaps represents the biggest source of bias in the fossil
record. First and foremost, organisms that contain hard parts have a
far greater chance of being represented in the fossil record than
organisms consisting of soft tissue only. As a result, animals with
bones or shells are overrepresented in the fossil record, and many
plants are only represented by pollen or spores that have hard walls.
Soft bodied organisms may form 30% to 100% of the biota, but most
fossil assemblages preserve none of this unseen diversity, which may
exclude groups such as fungi and entire animal phyla from the fossil
record. =June 2013 Many animals that moult, on the other hand,
are overrepresented, as one animal may leave multiple fossils due to
its discarded body parts. Among plants, wind-pollinated species
produce so much more pollen than animal-pollinated species, that the
former are much overrepresented relative to the latter.
Characteristics of the habitat
Most fossils form in conditions where material is deposited to the
bottom of water bodies. Especially shallow sea coasts produce large
amounts of fossils, so organisms living in such conditions have a much
higher chance of being preserved as fossils than organisms living in
non-depositing conditions. In continental environments, fossilization
is especially likely in small lakes that gradually fill in with
organic and inorganic material and especially in peat-accumulating
wetlands. The organisms of such habitats are therefore overrepresented
in the fossil record.
Mixing of fossils from different places
A sedimentary deposit may have experienced a mixing of
noncontemporaneous remains within single sedimentary units via
physical or biological processes; i.e. a deposit could be ripped up
and redeposited elsewhere, meaning that a deposit may contain a large
amount of fossils from another place (an allochthonous deposit, as
opposed to the usual autochthonous). Thus, a question that is often
asked of fossil deposits is to what extent does the fossil deposit
record the true biota that originally lived there? Many fossils are
obviously autochthonous, such as rooted fossils like crinoids, and
many fossils are intrisically obviously allocthonous, such as the
presence of photoautotrophic plankton in a benthic deposit that must
have sunk to be deposited. A fossil deposit may thus become biased
towards exotic species (i.e. species not endemic to that area) when
the sedimentology is dominated by gravity driven surges, such as
mudslides, or may become biased if there is very little endemic
organisms to be preserved. This is a particular problem in palynology.
Because population turnover rates of individual taxa are much less
than net rates of sediment accumulation, the biological remains of
successive, noncontemporaneous populations of organisms may be admixed
within a single bed, known as time-averaging. Because of the slow and
episodic nature of the geologic record, two apparently contemporaneous
fossils may have actually lived centuries, or even millennia, apart.
Moreover, the degree of time averaging in an assemblage may vary. The
degree varies on many factors, such as tissue type, the habitat, the
frequency of burial events and exhumation events, and the depth of
bioturbation within the sedimentary column relative to net sediment
accumulation rates. Like biases in spatial fidelity, there is a bias
towards organisms that can survive reworking events, such as shells.
An example of a more ideal deposit with respect to time-averaging bias
would be a volcanic ash deposit, which captures an entire biota caught
in the wrong place at the wrong time (e.g. the
Gaps in time series
The geological record is very discontinuous, and deposition is
episodic at all scales. At the largest scale, a sedimentological
high-stand period may mean that no deposition may occur for tens of
thousands of years and, in fact, erosion of the deposit may occur.
Such a hiatus is called an unconformity. Conversely, a catastrophic
event such as a mudslide may overrepresent a time period. At a shorter
scale, scouring processes such as the formation of ripples and dunes
and the passing of turbidity currents may cause layers to be removed.
Thus the fossil record is biased towards periods of greatest
sedimentation; periods of time that have less sedimentation are
consequently less well represented in the fossil record.
A related problem is the slow changes that occur in the depositional
environment of an area; a deposit may experience periods of poor
preservation to, for example, a lack of biomineralizing elements. This
causes the taphonomic or diagenetic obliteration of fossils, producing
gaps and condensation of the record.
Consistency in preservation over geologic time
Major shifts in intrinsic and extrinsic properties of organisms,
including morphology and behavior in relation to other organisms or
shifts in the global environment, can cause secular or long-term
cyclic changes in preservation (megabias).
Much of the incompleteness of the fossil record is due to the fact
that only a small amount of rock is ever exposed at the surface of the
Earth, and not even most of that has been explored. Our fossil record
relies on the small amount of exploration that has been done on this.
Unfortunately, paleontologists as humans can be very biased in their
methods of collection; a bias that must be identified. Potential
sources of bias include,
Search images: field experiments have shown that paleontologists
working on, say fossil clams are better at collecting clams than
anything else, because their search image has been shaped to bias them
in favour of clams.
Relative ease of extraction: fossils that are easy to obtain (such as
many phosphatic fossils that are easily extracted en masse by
dissolution in acid) are overabundant in the fossil record.
Taxonomic bias: fossils with easily discernible morphologies will be
easy to distinguish as separate species, and will thus have an
Preservation of biopolymers
Main article: Preservation of biopolymers
Although chitin exoskeletons of arthropods such as insects and
myriapods (but not trilobites, which are mineralized with calcium
carbonate, nor crustaceans, which are often mineralized with calcium
phosphate) are subject to decomposition, they often maintain shape
during permineralization, especially if they are already somewhat
The taphonomic pathways involved in relatively inert substances such
as calcite (and to a lesser extent bone) are relatively obvious, as
such body parts are stable and change little through time. However,
the preservation of "soft tissue" is more interesting, as it requires
more peculiar conditions. While usually only biomineralised material
survives fossilisation, the preservation of soft tissue is not as rare
as sometimes thought.
Both DNA and proteins are unstable, and rarely survive more than
hundreds of thousands of years before degrading. Polysaccharides
also have low preservation potential, unless they are highly
cross-linked; this interconnection is most common in structural
tissues, and renders them resistant to chemical decay. Such
tissues include wood (lignin), spores and pollen (sporopollenin), the
cuticles of plants (cutan) and animals, the cell walls of algae
(algaenan), and potentially the polysaccharide layer of some
lichens. This interconnectedness makes the chemicals
less prone to chemical decay, and also means they are a poorer source
of energy so less likely to be digested by scavenging organisms.
After being subjected to heat and pressure, these cross-linked organic
molecules typically "cook" and become kerogen or short (<17 C
atoms) aliphatic/aromatic carbon molecules. Other factors affect
the likelihood of preservation; for instance scleritisation renders
the jaws of polychaetes more readily preserved than the chemically
equivalent but non-sclerotised body cuticle.
It was thought that only tough, cuticle type soft tissue could be
preserved by Burgess Shale type preservation, but an increasing
number of organisms are being discovered that lack such cuticle, such
as the probable chordate
Pikaia and the shellless Odontogriphus.
It is a common misconception that anaerobic conditions are necessary
for the preservation of soft tissue; indeed much decay is mediated by
sulfate reducing bacteria which can only survive in anaerobic
conditions. Anoxia does, however, reduce the probability that
scavengers will disturb the dead organism, and the activity of other
organisms is undoubtedly one of the leading causes of soft-tissue
Plant cuticle is more prone to preservation if it contains cutan,
rather than cutin.
Plants and algae produce the most preservable compounds, which are
listed according to their preservation potential by Tegellaar (see
How complete fossils are was once thought to be a proxy for the energy
of the environment, with stormier waters leaving less articulated
carcasses. However, the dominant force actually seems to be predation,
with scavengers more likely than rough waters to break up a fresh
carcass before it is buried. Sediments cover smaller fossils
faster so they are likelier to be found fully articulated. However,
erosion also tends to destroy smaller fossils more easily.
Taphonomic processes allow researchers of multiple fields to identify
the past of natural and cultural objects. From the time of death or
burial until excavtion, taphonomy can aid in the understanding of past
environments. When studying the past it is important to gain
contextual information in order to have a solid understanding of the
data. Often these findings can be used to better understand cultural
or environmental shifts within the present day.
Trilobite type preservation
Bitter Springs type preservation
Burgess Shale type preservation
Doushantuo type preservation
Ediacaran type preservation
^ a b Lyman, R. Lee (2010-01-01). "What
Taphonomy Is, What it Isn't,
and Why Taphonomists Should Care about the Difference". Journal of
taphonomy. 8 (1): 1–16. ISSN 1696-0815.
^ Efremov, I. A. (1940). "Taphonomy: a new branch of paleontology".
Pan-American Geology. 74: 81–93. Archived from the original on
^ Martin, Ronald E. (1999) "1.1 The foundations of taphonomy"
Taphonomy: A Process Approach Cambridge University Press, Cambridge,
England, p. 1, ISBN 0-521-59833-8
^ "TAPHONOMY". personal.colby.edu. Retrieved 2017-05-03.
Taphonomy & Preservation". paleo.cortland.edu. Retrieved
^ Behrensmeyer, A. K; S. M Kidwell; R. A Gastaldo (2009), Taphonomy
^ Grotzinger, John P. (24 January 2014). "•Introduction to Special
Issue: Habitability, Taphonomy, and the Search for Organic Carbon on
Mars". Science. 343 (6169): 386–387. doi:10.1126/science.1249944.
PMID 24458635. Retrieved 2014-01-24.
^ Steele, Andrew; Beaty, David, eds. (September 26, 2006). "Final
report of the MEPAG Astrobiology Field Laboratory Science Steering
Group (AFL-SSG)". The Astrobiology Field Laboratory. U.S.A.: Mars
Exploration Program Analysis Group (MEPAG) – NASA. p. 72.
^ a b Grotzinger, John P. (January 24, 2014). "Introduction to Special
Issue – Habitability, Taphonomy, and the Search for Organic Carbon
on Mars". Science. 343 (6169): 386–387. doi:10.1126/science.1249944.
PMID 24458635. Retrieved January 24, 2014.
^ a b Various (January 24, 2014). "
Special Issue - Table of Contents -
Exploring Martian Habitability". Science. 343 (6169): 345–452.
Retrieved January 24, 2014. CS1 maint: Uses authors parameter
^ Various (January 24, 2014). "
Special Collection – Curiosity –
Exploring Martian Habitability". Science. Retrieved January 24,
2014. CS1 maint: Uses authors parameter (link)
^ Grotzinger, J.P. et al. (January 24, 2014). "A Habitable
Fluvio-Lacustrine Environment at Yellowknife Bay, Gale Crater, Mars".
Science. 343 (6169): 1242777. doi:10.1126/science.1242777.
PMID 24324272. Retrieved January 24, 2014. CS1 maint: Uses
authors parameter (link)
^ Passalacqua, Nicholas. "Introduction to Part VI: Forensic
^ admin (2011-12-08). "Forensic taphonomy". Crime Scene Investigator
(CSI) and forensics information. access-date= requires url=
^ "Front Matter". Manual of Forensic Taphonomy: i–xiv. 2013.
doi:10.1201/b15424-1. ISBN 978-1-4398-7841-5.
^ Fernandez Jalvo, Yolanda and Peter Andrews, “Methods in
Taphonomy” in Atlas of Taphonomic Identifications: 1001+ Images of
Fossil and Recent Mammal Bone Modification, ed. Eric Delson and Eric
J. Sargis Vertebrate
Paleobiology and Paleoanthropology Series (New
York, NY, American Museum of Natural History, 2016).
^ Rainsford C., and O'Connor T. 2016. "
Taphonomy and Contextual
Zooarchaeology in Urban Deposits at York, UK." Archaeological and
Anthropological Sciences 8 (2): 343–351.
^ Kidwell, S. M.; P. J Brenchley; D. Jablonski; D. H. Erwin; J. H.
Lipps (1996), "Evolution of the fossil record: thickness trends in
marine skeletal accumulations and their implications", Evolutionary
paleobiology: in honor of James W. Valentine: 290
^ "EBSCOhost Login". search.ebscohost.com. Retrieved 4 April
^ Briggs, D.E.G.; Kear, A.J. (1993), "Decay and preservation of
polychaetes; taphonomic thresholds in soft-bodied organisms",
Paleobiology, 19 (1): 107–135
^ a b c d e f g h i j Briggs, D.E.G. (1999), "Molecular taphonomy of
animal and plant cuticles: selective preservation and diagenesis",
Philosophical Transactions of the Royal Society B: Biological
Sciences, 354 (1379): 7–17, doi:10.1098/rstb.1999.0356,
^ Butterfield, N.J. (1990), "Organic preservation of non-mineralizing
organisms and the taphonomy of the Burgess Shale", Paleobiology, 16
(3): 272–286, JSTOR 2400788, (Registration required
^ Conway Morris, S. (2008), "A Redescription of a Rare Chordate,
Metaspriggina walcotti Simonetta and Insom, from the Burgess Shale
(Middle Cambrian), British Columbia, Canada", Journal of Paleontology,
82 (2): 424–430, doi:10.1666/06-130.1
^ Tegelaar, E.W.; De Leeuw, J.W.; Derenne, S.; Largeau, C. (1989), "A
reappraisal of kerogen formation", Geochim. Cosmochim. Acta, 53 (3):
^ Behrensmeyer, A. K.; Kidwell, S. M.; Gastaldo, R. A. (2000).
Taphonomy and Paleobiology". Paleobiology. 26 (4): 103–147.
^ "EBSCOhost Login". search.ebscohost.com. Retrieved 4 April
^ Lyman, R. Lee. Vertebrate taphonomy. Cambridge: Cambridge University
Emig, C. C. (2002). "Death: a key information in marine palaeoecology"
in Current topics on taphonomy and fossilization, Valencia". Col.lecio
Encontres. 5: 21–26.
Greenwood, D. R. (1991), "The taphonomy of plant macrofossils". In,
Donovan, S. K. (Ed.), The processes of fossilisation,
p. 141–169. Belhaven Press.
Lyman, R. L. (1994), Vertebrate Taphonomy. Cambridge University Press.
Shipman, P. (1981), Life history of a fossil: An introduction to
taphonomy and paleoecology. Harvard University Press.
Taylor, P. D.; Wilson, M. A. (2003). "Palaeoecology and evolution of
marine hard substrate communities" (PDF). Earth-Science Reviews. 62:
doi:10.1016/s0012-8252(02)00131-9. Archived from the original (PDF) on
The Shelf and Slope Experimental
Taphonomy Initiative is the first
long-term large-scale deployment and re-collection of organism remains
on the sea floor.
Journal of Taphonomy
Bioerosion Website at the College of Wooster
Comprehensive bioerosion bibliography compiled by Mark A. Wilson
Minerals and the Origins of Life (Robert Hazen, NASA) (video, 60m,
The 7th International Meeting on
Taphonomy and Fossilization (Taphos
2014) has been held at the Università degli Studi di Ferrara, Italy,
from 10 to 13 of September 2014.