Classification Principles

Trace fossils can be classified according to Linnaean taxonomy, ethology (inferred life history and
behaviour of trace producing organisms), or preservational type.

Linnaean classification involves the using of a binomial system similar to that employed for
body fossils and living species. Ichnotaxa include ichnogenera and ichnospecies, simply
used to define morphological differences. In many cases is difficult to prove a direct
relationship between trace producer and trace (e.g.. the trace produced by the
genus Lithophaga is called, according to its shape, Gastrochaenolites), therefore is desirable
an independent name for a particular trace morphology.

Ethological (Behavioural) classification is a mechanism of distinguishing groups of traces

on the basis of the behaviour of a group of organisms. The main types include:
o resting traces - cubichnia - created by active organisms while at rest or hiding from
prey;
o locomotion traces - repichnia - tracks and trails created while moving across the
sediment surface;
o dwelling traces - domichnia - three dimensional dwelling structures created by
burrowing;
 o grazing traces - pascichnia - horizontal feeding traces on the surface of the substrate;
o feeding traces - fodinichnia - three dimensional networks characterized by the
combined functions of deposit feeding and dwelling.
o traps and gardening traces - agrichnia - regular, patterned branching structures used
as traps for migrating meiofauna or as gardening systems where microbes might be
cultured for food.
o predation traces - praedichnia - are common on hard substrates, as round drill holes
in shells and shell damage by predators.
o equilibrium traces - equilibrichnia - are characterized by burrows that must be
constantly adjusted within the substrate due the agradation and degradation
processes.
o escape traces - fugichnia - occur as the animal flees to the new sea floor
o edifices built above the substrate - aedificichnia - are structures built of sediment,
more or less cemented by the architect
o structures made for breeding purposes - calichnia - are structures built for raising
larvae and juveniles.

Major categories of trace fossils according to behavioural classification and some illustrated
ichnogenera: Cruziana; 2. Anomoepus; 3. Cosmorhaphe; 4. Paleodictyon; 5. Phycosiphon; 6.
Zoophycos; 7. Thalassinoides; 8. Ophiomorpha; 9. Diplocraterion; 10. Gastrochaenolites; 11.
Asteriacites; Rusophycus (after Benton & Harper, 1997)

Preservational classification uses the relationship of the trace with the sedimentary surface.
According to this criterion the traces can be exogenic (on the sediment surface)
or endogenic (within the sediment itself). Traces can be found in full relief(with the
boundaries of circular burrows clearly defined within the enclosing sediment) or semirelief,
either on the upper surface (epirelief), or on the lower surface (hyporelief) of a sedimentary
layer.

Terminology of trace fossil preservation depending on the relationship to sediment horizons

(after Benton & Harper, 1997)
Trace fossils are usually identified using the combination of all the classification principles presented
above.

Types of Ichnofacies

The trace fossils have remained in appearance rather constant since Cambrian, even if their producers
might have been different. The trace fossil assemblages can be divided according the
palaeoenvironmental scheme into a number of ichnofacies named after a characteristic trace fossil. The
ichnofacies indicate a particular sedimentary facies and can be identified on the basis of its trace fossil
assemblage

The typical position of the major ichnofacies in marine and continental environments: Cr - Cruziana; G
- Glossifungites;
 N - Nereites; Ps - Psilonichnus; Sc - Scoyenia; Sk - Skolithos; Te - Teredolites; Tr - Tripanites; Z -
Zoophycos

(after Benton & Harper, 1997)

Loose- and Sedimentology /

Firmground
softground environment
Woodgrou Rockgrou
nd nd
Freshwat Freshwate Energ Bathymet Grainsi
Marine Marine
er r y ry ze

Psilonichn
- - Backshore Sand
us
Scoyenia
Rusophyco
Skolithos High Beach Sand
s?

Arenicolite Arenicolite
Event Shelf Sand silt
Glossifungit s? s
Teredolites Trypanites
es
Lagoon /
Fuersichnu Mediu Sand,
Cruziana
s? m silt
- shelf

Sand,
Nereites Event Slope to
mud
Mermia
abyssal
Zoophycos Low Mud

Scheme indicating relationships of ichnofacies with environment (after Bromley, 1996).

Nereites Ichnofacies
The Nereites Ichnofacies is recognized by the presence of meandering
pascichnia (Nereites, Neonereites and Helminthoide), spiral pascichnia (Spirorhaphe),
and agrichnia (Paleodictyon and Spirodesmos). Vertical burrows are almost entirely absent.

Nereites Ichnofacies viewed in pelagic carbonate ooze.

 (after Benton & Harper, 1997)

This ichnofacies indicates deep-water environments, including ocean floors and deep marine basins.
The trace fossils occur in muds deposited from suspension, and in the mudstones and siltstones of
distal turbidites.

Zoophycos Ichnofacies
The Zoophycos Ichnofacies is characterized by complex fodinichnia (Zoophycos, and sometimes other
deep traces such as Thalassinoides) in tiered arrangements.

(after Benton & Harper, 1997)

The ichnofacies occurs in a range of water depths between the abyssal zone and the shallow
continental shelf, in normal background conditions of sedimentation. The Nereites Ichnofacies may be
a matching association found at similar water depths during times of turbidite (event) deposition.

Cruziana Ichnofacies
The Cruziana Ichnofacies shows rich trace fossil diversity, with
horizontal repichnia (Cruziana and Aulichnites), cubichnia (Rusophycus, Asteriacites and Lockeia),
and vertical burrows.

(after Benton & Harper, 1997)

This ichnofacies represents mid and distal continental shelf situations, below normal wave base, but
may be affected by storm activity.

Skolithos Ichnofacies
The Skolithos Ichnofacies can be recognized by a low diversity of abundant
vertical domichniaburrows (Skolithos, Diplocraterion and Arenicolites), fodinichnia (Ophiomorpha),
and fugichnia.
 (after Benton & Harper, 1997)

All these traces typically indicate intertidal situations where the organisms have to be able to respond
rapidly in stressful conditions. The Skolithos Ichnofacies was at first seen as occurring only in the
intertidal zone, but it is also typical of other shifting sand environments, such as the tops of storm sand
sheets and the tops of turbidity flows.

Psilonichnus Ichnofacies
The Psilonichnus Ichnofacies shows a low diversity assemblage of small vertical burrows with basal
living chambers (Macanopsis), narrow sloping T-shaped and Y- shaped burrows (Psilonichnus), root
traces, and vertebrate footprints.

(after Benton & Harper, 1997)

This ichnofacies is typical of backshore, dune areas, and supratidal flats on the coast

Scoyenia Ichnofacies
The Scoyenia Ichnofacies is characterized by a low diversity trace fossil assemblage, mainly simple
horizontal fodinichnia (Scoyenia and Taenidium), with occasional vertical domichnia (Skolithos)
and repichnia produced by insects or freshwater shrimps (Cruziana, Isopodichnus).

(after Benton & Harper, 1997)

The traces are preserved in fluvial and lacustrine sediments, often in the silts and sands of redbed
sequences. Associated subaerial palaeosoils and aeolian sands may contain domichnia and repichnia of
insects, and dinosaur and other tetrapod foot-prints.
Glossifungites Ichnofacies
The Glossifungites Ichnofacies is characterized by domichnia (Glossifungites and Thalassinoides) and
sometimes plant root penetration structures. Other behavioural trace fossil types are rare.

(after Benton & Harper, 1997)

The traces occur in firm, but not lithified sediments, such as muds and silts in marine intertidal and
shallow subtidal zones. The firm grounds may develop in low energy situations such as salt marshes,
mud bars, of high intertidal flats, or in shallow marine environments where erosion has stripped off
superficial unconsolidated layers of sediment.

Trypanites Ichnofacies
The Trypanites Ichnofacies is characterized by domichnial borings of worms (Trypanites), bivalves
(Gostrochaenolites), barnacles (Rogerella) and sponges (Entobia).

(after Benton & Harper, 1997)

These are formed in shoreline rocks or in lithified limestone hard grounds on the seabed. Bioerosion
traces made by gastropods and echinoids are rarely preserved in ancient cases.

Teredolites Ichnofacies
The Teredolites Ichnofacies is identified by the presence of borings in wood (especially Teredolites),
especially those produced by marine bivalves such as the modern ship worm, Teredo.

(after Benton & Harper, 1997)