Trace fossil

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Chirotherium footprints in a Triassicsandstone.

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Protichnites tracksway from theCambrian, Blackberry Hill, central Wisconsin.

Trace fossils, also called ichnofossils (sg. /knofsl/; Greek: ikhnos "trace, track"),
are geological records of biological activity. Trace fossilsmay be impressions made on 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 is called ichnology, which is divided into paleoichnology, or the study of trace fossils,
and neoichnology, the study of modern traces. This science is challenging, as most traces reflect the behaviour
not the biological affinity of their makers. As such, trace fossils are categorised into form genera, based
upon their appearance and the implied behaviour of their makers.
Contents
[hide]

1 Occurrence

2 Classification

3 Information provided by ichnofossils

3.1 Paleoecology

3.2 Paleoenvironment

3.3 Stratigraphic correlation

4 Ichnofacies

5 Inherent bias

6 Evolution

7 Common ichnogenera

8 Other notable trace fossils

9 Confusion with other types of fossils

10 See also

11 References

12 Further reading

13 External links

Occurrence[edit]

Cross-section of mammoth footprints at The Mammoth Site, Hot Springs, South Dakota.

Traces are better known in their fossilised form than in modern sediments. [1] This makes it difficult to interpret
some fossils by comparing them with modern traces, even though they may be extant or even common. [1] The
main difficulties in accessing extant burrows stem from finding them in consolidated sediment, and being able
to access those formed in deeper water.
Trace fossils are best preserved in sandstones; [1] the grain size and depositional facies both contributing to the
better preservation. They may also be found in shales and limestones. [1]

Classification[edit]
Main article: Trace fossil classification
Trace fossils are generally difficult or impossible to assign to a specific maker. Only in very rare occasions are
the makers found in association with their tracks. Further, entirely different organisms may produce identical
tracks. Therefore conventional taxonomy is not applicable, and a comprehensive form taxonomy has been
erected. At the highest level of the classification, five behavioral modes are recognized: [1]

Domichnia, dwelling structures reflecting the life position of the organism that created it.

Fodinichnia, three-dimensional structures left by animals which eat their way through sediment, such
as deposit feeders;

Pascichnia, feeding traces left by grazers on the surface of a soft sediment or a mineral substrate;

Cubichnia, resting traces, in the form of an impression left by an organism on a soft sediment;

Repichnia, surface traces of creeping and crawling.

Fossils are further classified into form genera, a few of which are even subdivided to a "species" level.
Classification is based on shape, form, and implied behavioural mode.

Information provided by ichnofossils[edit]

Mesolimulus walchi fossil and track, a rare example of tracks and the creature that made them fossilized together

Because identical fossils can be created by a range of different organisms, trace fossils can only reliably inform
us of two things: the consistency of the sediment at the time of its deposition, and the energy level of
the depositional environment.[2] Attempts to deduce such traits as whether a deposit is marine or non-marine
have been made, but shown to be unreliable.[2]

Paleoecology[edit]
Trace fossils provide us with indirect evidence of life in the past, such as the footprints, tracks, burrows,
borings, and feces left behind by animals, rather than the preserved remains of the body of the actual animal
itself. Unlike most other fossils, which are produced only after the death of the organism concerned, trace
fossils provide us with a record of the activity of an organism during its lifetime.
Trace fossils are formed by organisms performing the functions of their everyday life, such as walking,
crawling, burrowing, boring, or feeding. Tetrapod footprints, worm trails and the burrows made
by clams andarthropods are all trace fossils.
Perhaps the most spectacular trace fossils are the huge, three-toed footprints produced by dinosaurs and
related archosaurs. These imprints give scientists clues as to how these animals lived. Although the skeletons
of dinosaurs can be reconstructed, only their fossilized footprints can determine exactly how they stood and
walked. Such tracks can tell much about the gait of the animal which made them, what its stride was, and
whether or not the front limbs touched the ground.
However, most trace fossils are rather less conspicuous, such as the trails made by segmented
worms or nematodes. Some of these worm castings are the only fossil record we have of these soft-bodied
creatures.

Paleoenvironment[edit]

Eubrontes, a dinosaur footprint in the Lower Jurassic Moenave Formation at the St. George Dinosaur Discovery Site at
Johnson Farm, southwestern Utah.

Fossil footprints made by tetrapod vertebrates are difficult to identify to a particular species of animal, but they
can provide valuable information such as the speed, weight, and behavior of the organism that made them.
Such trace fossils are formed when amphibians, reptiles, mammals or birds walked across soft (probably wet)
mud or sand which later hardened sufficiently to retain the impressions before the next layer of sediment was
deposited. Some fossils can even provide details of how wet the sand was when they were being produced,
and hence allow estimation of paleo-wind directions.[3]
Assemblages of trace fossils occur at certain water depths, [1] and can also reflect the salinity and turbidity of the
water column.

Stratigraphic correlation[edit]
Some trace fossils can be used as local index fossils, to date the rocks in which they are found, such as the
burrow Arenicolites franconicus which occurs only in a 4 cm (1.6") layer of the Triassic Muschelkalk epoch,
throughout wide areas in southern Germany.[4]
The base of the Cambrian period is defined by the first appearance of the trace fossil Treptichnus pedum.[5]
Trace fossils have a further utility as many appear before the organism thought to create them, extending their
stratigraphic range.[6]

Ichnofacies[edit]

Main article: Ichnofacies

Trace fossil assemblages are far from random; the range of fossils recorded in association is constrained by
the environment in which the trace-making organisms dwelt.[1] Palaeontologist Adolf Seilacher pioneered the
concept of ichnofacies, whereby the state of a sedimentary system at its time of deposition could be implied by
noting the fossils in association with one another.[1]

Inherent bias[edit]

Diagram showing how dinosaur footprints are preserved in different deposits

Most trace fossils are known from marine deposits. [7] Essentially, there are two types of traces, either exogenic
ones, which are made on the surface of the sediment (such as tracks) or endogenic ones, which are made
within the layers of sediment (such as burrows).
Surface trails on sediment in shallow marine environments stand less chance of fossilization because they are
subjected to wave and current action. Conditions in quiet, deep-water environments tend to be more favorable
for preserving fine trace structures.
Most trace fossils are usually readily identified by reference to similar phenomena in modern environments.
However, the structures made by organisms in recent sediment have only been studied in a limited range of
environments, mostly in coastal areas, including tidal flats.[citation needed]

Evolution[edit]

Climactichnites, probably trackways from a slug-like animal, from the Cambrian, Blackberry Hill, central Wisconsin. Ruler in
background is 45cm (18") long.

The earliest complex trace fossils, not including microbial traces such as stromatolites, date
to2,000 to 1,800 million years ago. This is far too early for them to have an animal origin, and they are thought
to have been formed by amoedae.[8] Putative "burrows" dating as far back as 1,100 million years may have
been made by animals which fed on the undersides of microbial mats, which would have shielded them from a
chemically unpleasant ocean;[9] however their uneven width and tapering ends make a biological origin so
difficult to defend[10] that even the original author no longer believes they are authentic. [11]
The first evidence of burrowing which is widely accepted dates to the Ediacaran (Vendian) period,
around 560 million years ago[verification needed]. During this period the traces and burrows basically are horizontal on
or just below the seafloor surface. Such traces must have been made by motile organisms with heads, which
would probably have been bilateran animals.[12] The trace observed imply simple behaviour, and point to
organisms feeding above the surface and burrowing for protection from predators. [13] Contrary to widely
circulated opinion that Ediacaran burrows are only horizontal the vertical burrows Skolithos are also known.
[14]

The producers of burrows Skolithos declinatus from the Vendian (Ediacaran) beds in Russia with

date555.3 million years ago have not been found, they might have been filter feeders subsisting on the
nutrients from the suspension. The density of these burrows is up to 245 burrows/dm 2.[15] Some Ediacaran trace
fossils have been found directly associated with an body fossils. Yorgia and Dickinsoniaare often found at the
end of long pathways of trace fossils matching their shape. [16] The feeding was performed in a mechanical way,
supposedly the ventral side of body these organisms was covered with cilia.[17] The
potential mollusc related Kimberella is associated with scratch marks, perhaps formed by a radula,[18] further
traces from 555 million years ago appear to imply active crawling or burrowing activity.[19]
As the Cambrian got underway, new forms of trace fossil appeared, including vertical burrows
(e.g. Diplocraterion) and traces normally attributed to arthropods.[20] These represent a widening of the
behavioural repertoire,[21] both in terms of abundance and complexity.[22]
Trace fossils are a particularly significant source of data from this period because they represent a data source
that is not directly connected to the presence of easily fossilized hard parts, which are rare during the
Cambrian. Whilst exact assignment of trace fossils to their makers is difficult, the trace fossil record seems to
indicate that at the very least, large, bottom-dwelling, bilaterally symmetrical organisms were rapidly
diversifying during the early Cambrian.[23]
Further, less rapid[verification needed] diversification occurred since,[verification needed] and many traces have been
converged upon independently by unrelated groups of organisms. [1]
Trace fossils also provide our earliest evidence of animal life on land. The earliest arthropod trackways date to
the Cambro-Ordovician,[24] and trackways from the Ordovician Tumblagooda sandstone allow the behaviour of

these organisms to be determined.[3] The enigmatic trace fossil Climactichnites may represent an earlier still
terrestrial trace, perhaps made by a slug-like organism. [verification needed]

Common ichnogenera[edit]

Petroxestes borings in a hardground from the Upper Ordovician of southern Ohio.

Rusophycus trace fossil from the Ordovician of southern Ohio. Scale bar is 10 mm.

Skolithos trace fossil. Scale bar is 10 mm.

Thalassinoides, burrows produced by crustaceans, from the Middle Jurassic,Makhtesh Qatan, southern Israel.

Trypanites borings in an UpperOrdovician hardground from northern Kentucky. The borings are filled with
diagenetic dolomite (yellowish). Note that the boring on the far right cuts through a shell in the matrix.

Asteriacites is the name given to the five-rayed fossils found in rocks and they record the resting place
of starfish on the sea floor. Asteriacites are found in European and American rocks, from
the Ordovician period onwards, and are numerous in rocks from the Jurassic period of Germany.

Chondrites (not to be confused with stony meteorites of the same name) are small branching burrows
of the same diameter, which superficially resemble the roots of a plant. The most likely candidate for
having constructed these burrows is a nematode (roundworm). Chondrites are found in marine sediments
from the Cambrian period of the Paleozoic onwards. They are especially common in sediments which were
deposited in reduced-oxygen environments.

Climactichnites is the name given to surface trails and burrows that consist of a series of chevronshaped raised cross bars that are usually flanked on either side by a parallel ridge. They somewhat
resemble tire tracks, and are larger (typically about four inches wide) than most of the other trace fossils
made by invertebrates. The trails were produced on sandy tidal flats during Cambrian time. While the
identity of the animal is still conjectural, it may have been a large slug-like animal - its trails produced as it
crawled over and processed the wet sand to obtain food. [25][26]

Cruziana are excavation trace marks made on the sea floor which have a two-lobed structure with a
central groove. The lobes are covered with scratch marks made by the legs of the excavating organism,
usually a trilobite or allied arthropod. Cruziana are most common in marine sediments formed during
the Paleozoic era, particularly in rocks from the Cambrian and Ordovician periods. Over 30 ichnospecies
of Cruziana have been identified. See also Isopodichnus.

Entobia is a boring produced by endolithic clionaid sponges consisting of galleries excavated in a

carbonate substrate; often has swollen chambers with connecting canals.

Gastrochaenolites are clavate (club-shaped) borings also produced in calcareous hard substrates,
usually by bivalves.

Petroxestes is a shallow groove boring produced by mytilacean bivalves in carbonate hard substrates.

Protichnites consists of two rows of tracks and a linear depression between the two rows. The tracks
are believed to have been made by the walking appendages of arthropods. The linear depression is
thought to be the result of a dragging tail. The structures bearing this name were typically made on the
tidal flats of Paleozoic seas, but similar ones extend into the Cenozoic.

Rhizocorallium is a type of burrow, the inclination of which is typically within 10 of the bedding planes
of the sediment. These burrows can be very large, over a meter long in sediments that show good
preservation, e.g. Jurassic rocks of the Yorkshire Coast (eastern United Kingdom), but the width is usually
only up to 2 cm, restricted by the size of the organisms producing it. It is thought that they represent
fodinichnia as the animal (probably a nematode) scoured the sediment for food.

Rogerella is a small pouch-shaped boring with a slit-like aperture currently produced

by acrothoracican barnacles.

Rusophycus are bilobed "resting traces" associated with trilobites and other arthropods such as
horseshoe crabs.

Skolithos: One well-known occurrence of Cambrian trace fossils from this period is the famous 'Pipe
Rock' of northwest Scotland. The 'pipes' that give the rock its name are closely packed straight tubeswhich were presumably made by some kind of worm-like organism. The name given to this type of tube or
burrow is Skolithos, which may be 30 cm (12") in length and between 2 to 4 cm (0.8 to 1.6") in diameter.
Such traces are known worldwide from sands and sandstones deposited in shallow water environments,
from the Cambrian period (542488 Ma) onwards.

Thalassinoides are burrows which occur parallel to the bedding plane of the rock and are extremely
abundant in rocks, worldwide, from the Jurassicperiod onwards. They are repeatedly branched, with a
slight swelling present at the junctions of the tubes. The burrows are cylindrical and vary from 2 to 5 cm
(0.8" to 2") in diameter. Thalassinoides sometimes contain scratch marks, droppings or the bodily remains
of the crustaceans which made them.

Teichichnus has a distinctive form produced by the stacking of thin 'tongues' of sediment, atop one
another. They are again believed to be fodinichnia, with the organism adopting the habit of retracing the
same route through varying heights of the sediment, which would allow it to avoid going over the same
area. These 'tongues' are often quite sinuous, reflecting perhaps a more nutrient-poor environment in
which the feeding animals had to cover a greater area of sediment, in order to acquire sufficient
nourishment.

Trypanites are elongated cylindrical borings in calcareous substrates such as shells,

carbonate hardgrounds and limestones. Usually produced by worms of various types and sipunculids.

Other notable trace fossils[edit]

Less ambiguous than the above ichnogenera, are the traces left behind by invertebrates such
as Hibbertopterus, a giant "sea scorpion" or eurypterid of the early Paleozoic era. This
marine arthropod produced a spectacular hibbertopteroid track preserved in Scotland. [27]
Bioerosion through time has produced a magnificent record of borings, gnawings, scratchings and scrapings on
hard substrates. These trace fossils are usually divided into macroborings [28] and microborings.[29] Bioerosion
intensity and diversity is punctuated by two events. One is called the Ordovician Bioerosion Revolution (see
Wilson & Palmer, 2006) and the other was in the Jurassic. [30] For a comprehensive bibliography of the
bioerosion literature, please see the External links below.
The oldest types of tetrapod tail-and-foot prints date back to the latter Devonian period.
These vertebrate impressions have been found in Ireland,Scotland, Pennsylvania, and Australia.
Important human trace fossils are the Laetoli (Tanzania) footprints, imprinted in volcanic ash 3.7 Ma (million
years ago) -- probably by an earlyAustralopithecus.[31]

Confusion with other types of fossils[edit]

Asteriacites (sea star trace fossil) from the Devonian of northeastern Ohio. It appears at first to be an external mold of the
body, but the sediment piled between the rays shows that it is a burrow.

Trace fossils are not body casts. The Ediacara biota, for instance, primarily comprises the casts of organisms in
sediment. Similarly, a footprint is not a simple replica of the sole of the foot, and the resting trace of a seastar
has different details than an impression of a seastar.
Early paleobotanists misidentified a wide variety of structures they found on the bedding planes of sedimentary
rocks as fucoids (Fucales, a kind ofbrown algae or seaweed). However, even during the earliest decades of the
study of ichnology, some fossils were recognized as animal footprints and burrows. Studies in the 1880s by A.
G. Nathorst and Joseph F. James comparing 'fucoids' to modern traces made it increasingly clear that most of
the specimens identified as fossil fucoids were animal trails and burrows. True fossil fucoids are quite rare.
Pseudofossils, which are not true fossils, should also not be confused with ichnofossils, which are true
indications of prehistoric life.

Numerous borings in a Cretaceous cobble, Faringdon, England; see Wilson (1986).

Sponge borings (Entobia) and encrusters on a modern bivalve shell, North Carolina.

Helminthopsis ichnosp.; a trace fossil from the Logan Formation (Lower Carboniferous) of Wooster, Ohio.

Gigandipus, a dinosaur footprint in the Lower Jurassic Moenave Formation at the St. George Dinosaur Discovery
Site at Johnson Farm, southwestern Utah.

Lockeia from the Dakota Formation(Upper Cretaceous).

Lockeia from the Chagrin Shale (Upper Devonian) of northeastern Ohio. This is an example of the trace fossil
ethological group Fugichnia.

Gnathichnus pentax echinoid trace fossil on an oyster from theCenomanian of Hamakhtesh Hagadol, southern
Israel.

See also[edit]
Wikimedia Commons has
media related to Trace fossils.

Bioerosion

Bird ichnology

Fossil egg

Ichnite - fossilised footprints

Ichnofacies

Index fossil

Trace fossil classification

Way up structure

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Further reading[edit]

Bromley, R.G., 1970. Borings as trace fossils and Entobia cretacea Portlock as an example, p. 49-90.
In: Crimes, T.P. and Harper, J.C. (eds.), Trace Fossils. Geological Journal Special Issue 3.

Bromley, R.G., 2004. A stratigraphy of marine bioerosion. In: The application of ichnology to
palaeoenvironmental and stratigraphic analysis. (Ed. D. McIlroy), Geological Society of London Special
Publications 228:455-481.

Palmer, T.J., 1982. Cambrian to Cretaceous changes in hardground communities. Lethaia 15:309-323.

Wilson, M.A., 1986. Coelobites and spatial refuges in a Lower Cretaceous cobble-dwelling hardground
fauna. Palaeontology 29:691-703.

Wilson, M.A. and Palmer, T.J., 2006. Patterns and processes in the Ordovician Bioerosion Revolution.
Ichnos 13: 109-112.[2]

Yochelson, E.L. and Fedonkin, M.A., 1993. Paleobiology of Climactichnites, and Enigmatic Late
Cambrian Fossil. Smithsonian Contributions to Paleobiology 74:1-74.

External links[edit]

Encyclopaedia-style article about trace fossils

Ichnogenus images

Bioerosion Website at The College of Wooster

Comprehensive bioerosion bibliography compiled by Mark A. Wilson