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TRACE FOSSILS
P J Orr, University College Dublin, Dublin, Ireland ichnofabrics has made a fundamental and growing
ß 2005, Elsevier Ltd. All Rights Reserved. contribution to our understanding of the evolutionary
palaeoecology of the Earth’s biosphere. Furthermore,
bioturbation impacts on the physical and chemical
Introduction properties of sediments, including their diagenesis.
Key terms employed in the taxonomic, preservational,
Trace fossils and ichnofabrics offer an alternative and ethological classification of trace fossils are de-
source of data on the ecology of any palaeoecosystem fined; how trace fossils and ichnofabrics are used in
to that provided by the body fossil record. Different palaeoenvironmental and community reconstructions
preservational biases apply to the trace and body is discussed.
fossil records. ‘Soft-bodied’ organisms, i.e., those ‘Trace fossil’ is used herein to describe any discrete
lacking biomineralized tissues, can produce trace structure produced by the reworking of sediment
fossils, yet their fossilization potential is minimal. or the bioerosion of lithic (rock) or xylic (wood)
Most trace fossils are emplaced into unconsolidated substrates by infauna or epifauna. Eggs and sediment-
sediment, cannot survive reworking, and are thus ary structures produced by physical processes do
autochthonous (in marked contrast to the vast major- not represent trace fossils. Some authorities include
ity of body fossils). The study of trace fossils and stromatolites within this term, on which basis
 TRACE FOSSILS 521

Figure 1 Wrinkle structures in siliciclastic lithologies attributed

to the former presence of a microbial mat. Image courtesy of Figure 2 Style of burrowing reflects substrate consistency.
Séan Burke. (A) Intrusive burrowing typical of near-surface soupgrounds.
(B) Compression burrowing generates an open burrow and no
‘suspect microbial structures’ (e.g., wrinkle structures spoils. (C) Excavation, yielding an open burrow and advected
spoil. (D) Excavation, producing a backfilled burrow. In the latter
(Figure 1), roll-up structures, and elephant skin tex- case, the active infill, i.e., by the producer, may be structureless,
ture) would be included; these are attributed to the but is often sorted; examples include separation into a distinct
presence, including plastic deformation, of micro- core and marginal or (as here) meniscate structures. The geom-
bially bound crusts or veneers on the top of the sedi- etry of individual menisci can vary from shallow and saucer-like
ment column. An ichnofabric, all aspects of the to sharply pointed cones; successive menisci are often defined
by alternations in their colour and/or composition.
sedimentary texture that result from biogenic rework-
ing, includes any remnant sedimentary structures,
trace fossils, or indistinct structures produced by fossils. Some higher rank categories (ichnofamilies),
macrofauna (‘sediment churning’, ‘burrow mottling’, or informal groupings, have been established, but are
and biodeformational structures) that cannot be ac- not used widely. As a sedimentary structure, the
corded formal taxonomic status. The poor definition detailed morphology of a trace fossil reflects both
of the latter usually reflects their emplacement the producer’s behaviour and the properties of the
in sediments with a high pore water content and low host sediment; characteristics of the latter (such as
sediment shear strength (soupgrounds); it may be water content or grain size) can vary between beds,
compounded by the employment of a different bur- or even the laminae of a bed (Figures 3A and 3B). The
rowing strategy (Figure 2). Intrusive burrowing in abundance, probably surfeit, of ichnotaxa, particu-
uncompacted sediment involves simple deflection larly ichnospecies, in the geological literature has its
of the sediment around the body; the sediment col- origin in the middle to late nineteenth century, when
lapses behind the body to occlude any void immedi- many trace fossils were interpreted as body fossils;
ately (sediment swimming). In more compacted every subtle variation in morphology was therefore
sediments (softgrounds and firmgrounds), compres- considered to be significant, and the structure worthy
sion burrowing and, especially, excavation (accom- of separate classification. Further, and more easily
panied by either advection or backfilling) produce resolved by comprehensive study, the morphology of
more complex, open, and actively infilled structures. a trace fossil in partim, and thus the ichnotaxobases
Cryptobioturbation, resulting from the activities of (the features used to classify it), may vary depending
meiofauna and microfauna, may obliterate any pri- on the orientation in which it is observed (Figures 3B
mary sedimentary depositional fabric and homogen- and 3C).
ize sediment without producing either discrete trace
fossils or biodeformational structures. Identifying the producer Trace fossils are very rarely
preserved in association with their producer; if they
are, each should be named separately. The identifica-
Taxonomy, Preservation, tion of the causative organism is not a prerequisite
and Ethology for the naming of a trace fossil. One organism may
produce more than one trace fossil depending on its
Taxonomy
behaviour (Figure 4A). A structure may also be pro-
Trace fossils are classified using ichnogenus and ich- duced by different coexisting organisms (Figure 4B),
nospecies; the prefix ‘ichno’ distinguishes these or modified later by a different organism. Poten-
ichnotaxa from either living organisms or body tial producers may be constrained (usually at the
522 TRACE FOSSILS

Figure 3 Key concepts in the study of trace fossils 1. (A) Preservational variants of the same ichnotaxon, reflecting emplacement in
sediments of different consistency. (B) Different views of the same ichnotaxon reflect slight differences in the level of burrowing
relative to the clay–sand interface. From Ekdale AA, Bromley RG, and Pemberton SG (1984) Ichnology. The Use of Trace Fossils in
Sedimentology and Stratigraphy. SEPM Short Course No. 15. Tulsa, OK: Society of Economic Paleontologists and Mineralogists. (C) A slight
angular difference between the plane of bedding/weathering and that containing the trace fossil results in different two-dimensional
views of the trace fossil (1–3). Scale bars, 5 mm.

Figure 4 Key concepts in the study of trace fossils 2. (A) Open J-shaped dwelling burrow and excavated pelleted sand (1), trackway
(2), feeding structure (3), and faecal pellets (4) produced by the fiddler crab, Uca. (B) Plan view of open burrow network produced by a
crab, lobster, and fish. (C) Examples of Rusophycus produced by a polychaete, snail, trilobite, and notostracan. From Ekdale AA,
Bromley RG, and Pemberton SG (1984) Ichnology. The Use of Trace Fossils in Sedimentology and Stratigraphy. SEPM Short Course No. 15.
Tulsa, OK: Society of Economic Paleontologists and Mineralogists.

resolution of phylum or class) by the morphology fossil may be produced by a variety of animals,
and inferred function of the trace fossil (e.g., ‘arthro- precluding extrapolation of the producer’s identity
pod-produced trackway’). The body fossil content between case studies. The resting trace Rusophycus,
of the same, or surrounding, lithologies may be examples of which in Palaeozoic marine sediments
suggestive; for example, the breadth of a trackway are often attributed to trilobites (see Fossil Inverte-
may correspond to that of one, but not other, poten- brates: Trilobites), also occurs in Mesozoic strata
tial candidates. The potential producers of trace (after trilobites became extinct) and in sediments
fossils, however, include organisms with minimal from (non-marine) environments that were never
preservation potential. Furthermore, the same trace colonized by trilobites (Figure 4C).
 TRACE FOSSILS 523

Preservation substrate during emplacement of exogenic traces

may depress underlying sediment; in finely laminated
Exogenic trace fossils are emplaced at the sediment–
lithologies, splitting a short vertical distance below
water or sediment–air interface, and endogenic traces
the original interface may reveal such undertracks, the
within the substrate; intergenic trace fossils are those
resolution of which is often poorer than that of the
emplaced endogenously at the interface between two
corresponding exogenic expression; furthermore,
beds (Figure 5A). Exogenic trace fossils have limited
only those parts of the trace fossil where loading
fossilization potential; they must be covered and thus
was greater may be expressed as undertracks. Endo-
cast, not eroded, during later deposition. Splitting at
the original interface will yield both epirelief and genic trace fossils observed in three dimensions are in
full relief; those exposed on bedding-parallel surfaces
hyporelief views of the structure. Loading of the

Figure 5 Key concepts in the study of trace fossils 3. (A) Preservational classification of trace fossils. Adapted from Ekdale AA,
Bromley RG, and Pemberton SG (1984) Ichnology. The Use of Trace Fossils in Sedimentology and Stratigraphy. SEPM Short Course No. 15. Tulsa,
OK: Society of Economic Paleontologists and Mineralogists. (B) The intergenic trace fossil is exposed in semirelief (positive hyporelief)
on the sole of a bed of sandstone following preferential weathering of the underlying finer grained bed. (C) Positive epirelief view of
Lophoctenium showing differential weathering of alternate menisci. (D) Polished horizontal cross-section through an actively infilled
burrow comprising a thick marginal wall structure and meniscate core. (E) Formation of predepositional and postdepositional trace
fossils on the sole of an event bed. (F) Although the faecal pellets are the same colour and sourced from the host lithology, the light-
coloured sediment in the interstices between them indicates that the trace fossil is secondary. Scale bars, 5 mm.
524 TRACE FOSSILS

are in semirelief; hyporelief and epirelief are used critical conditions, erosion closely followed by depos-
when the lower or upper surface, respectively, is that ition, are often met during deposition of event beds,
exposed. The orientation of specimens in the field such as turbidites or storm deposits.
should be determined at the time of their collection Postdepositional intergenic burrows may be em-
using independent criteria provided by sedimentary placed later onto a surface containing predeposi-
structures. The prefixes positive and negative de- tional secondary casts (Figure 5E). As well as the
scribe the relief of the trace fossil relative to the host difference in their relative age, each suite may re-
lithology. Full relief, and particularly semirelief, views flect different environmental conditions. Only the
can result from planes of splitting deflecting to follow lower surface of a predepositional endogenic burrow
the external surface of, rather than continuing across, system will be preserved; its infill will comprise the
a trace fossil. Semirelief views of intergenic structures host lithology. Active infill of a postdepositional
are most common when successive lithologies have a burrow may include modification of the host lith-
different resistance to weathering (Figure 5B). Parts ology (see above); in vertical cross-section, the out-
of an individual structure may weather differently; in line of the burrow (whether infilled actively or, later,
the example in Figure 5C, alternate chevron-shaped passively) will be complete.
menisci are either more sand-rich or mud-rich than,
and thus have weathered positive and negative with Primary and secondary trace fossils Primary trace
respect to, the host lithology. Localized differences in fossils represent either reworking of the host lithology
the sedimentary fabric that result from bioturbation in situ (but not necessarily at the time of its depos-
can be exacerbated during diagenesis. Trace fossils ition), or the piping downwards of sediment of iden-
are often sites for the precipitation of early diagenetic tical composition from one layer into another. Their
minerals; this may be encouraged by the presence of identification relies on the properties of the host lith-
mucus secreted during burrowing as an aid to loco- ology being altered in the process (see above). Sec-
motion, excretion, or to maintain an open structure. ondary trace fossils originate when sediment with
This can impact on economically important variables different properties is ‘piped’ (usually downwards)
such as sediment texture, organic content, porosity, into the host lithology; the evidence can be subtle
and permeability. (see example in Figure 5F).
An open structure connected to either the air or
Ethology
water column may be later infilled gravitationally
(passive infill) by sediment of a different composition A behavioural, or ethological, classification of trace
and/or grain size. Active infill of a structure is by its fossils is presented in Figure 6. Few trace fossils
progenitor. Modifications resulting from the rework- represent just one activity, and classification is thus
ing of sediment, for example during its ingestion on the basis of what is considered to be the dominant,
and passage through the gut of an animal, include most significant, behaviour. Cubichnia, temporary
(1) excretion as pellets or a faecal string; (2) the resting traces, are usually shallow, exogenic excav-
exclusion or selection of grains by shape, size, or ations (Figure 7A). Repichnia, locomotion traces, in-
type; and (3) the reorientation of grains to produce a clude both endogenic continuous burrows and
localized sedimentary fabric (Figure 5D). exogenic structures. The latter may be continuous
(trails), or comprise a series of discrete sequential
Predepositional and postdepositional trace fossils footfalls (tracks or imprints) made by an appendage
Postdepositional trace fossils are emplaced after the or limb (trackways) (Figure 7B). Pascichnia combine
deposition of a bed; the term ‘predepositional’ is used continuous locomotion parallel to bedding with feed-
for trace fossils, now preserved on the sole of a bed, ing; emphasis is on the systematic extraction of food
that were emplaced on the surface of, or within, the resources within an area (cf., strip mining). This is
underlying bed. Thus, while an exogenic trackway achieved by phobotactic (the avoidance of crossing
in epirelief view is postdepositional, the secondary previously formed parts of the structure), combined
cast produced during deposition of the next bed, with strophotactic (episodic or periodic 180 turns)
and exposed subsequently in hyporelief on the sole and/or thigmotactic (tendency to keep close to a pre-
thereof, is predepositional. For exogenic trace fossils, viously formed part of the structure), behaviour pat-
the process requires there to be no erosion of the terns (Figures 7C and 7D). Trails in negative epirelief
existing sediment. More extensive erosion, however, with similar geometries on the surfaces of modern
may expose endogenic open burrow systems, or wash ocean floors are exogenic structures with limited
away the active infill of burrows whilst preserving fossilization potential; at least, the vast majority
their outlines; infill of the voids created generates of fossil pascichnia were actively filled endogenic
predepositional secondary casts (Figure 5E). The burrow systems. Agrichnia are endogenic burrow
 TRACE FOSSILS 525

Figure 6 An ethological classification of trace fossils. Adapted from Ekdale AA, Bromley RG, and Pemberton SG (1984) Ichnology. The
Use of Trace Fossils in Sedimentology and Stratigraphy. SEPM Short Course No. 15. Tulsa, OK: Society of Economic Paleontologists and
Mineralogists.

systems that were maintained as open structures, and identified; some examples of the fossilized cases of
are therefore almost always preserved in positive caddis larvae contain pupae (Figure 7K). Domichnia
hyporelief as predepositional secondary casts (Figures in soft sediments often exhibit passive infill from
5E and 7E). Their characteristic, complex geometries the overlying layer and a thick marginal lining (to
in plan view include fish scale-like, polygonal, and prevent collapse of the open burrow system);
planispiral patterns. Spiralling patterns maintain a examples include the pelleted walls typical of Ophio-
constant spacing between successive whorls, i.e., morpha. A domichnion in a firmground may have a
they do not exhibit thigmotactic behaviour. Many bioglyph on its internal surface (Figure 7L). A ‘spreite’
agrichnia are essentially two-dimensional, contained (plural spreiten) is produced by shifting the position
within a single plane parallel to the sediment–water of a U-shaped vertical burrow in order to maintain its
interface; three-dimensional structures comprise base a constant distance below the sediment–water
several such levels, separated from each other verti- interface. The spreite, which marks the former pos-
cally and connected by a more steeply inclined con- ition of the basal part of the burrow, may be pro-
tinuous open burrow. Agrichnia possessed single or trusive (occurring inside the limbs of the open
multiple vertical connections to the overlying water burrow system in response to sediment being re-
column. As with domichnia (see below), the introduc- moved; Figure 7I), and/or retrusive (occurring below
tion of oxygenated seawater to depth within the the active burrow in response to sediment aggrada-
sediment could have reduced the idealized vertical tion). Equilibrichnia, formed by the regular, incre-
stratification of electron acceptors to a patchwork mental shift upwards of burrows in response to the
of biogeochemical microenvironments (Figure 8). semicontinuous accretion of sediment, are considered
Their exact function is unknown, but agrichnia may here as a variety of domichnion. Rapid upward
represent traps or ‘microbial gardens’; microbes (attempted) escape, for example following burial by
may be cultured on, and harvested from, the walls. sediment, results in poorly structured escape traces,
Fodinichnia combine semistationary behaviour with fugichnia (fugichnion). Unequivocal examples of
deposit feeding. Geometries are highly variable and praedichnia, trace fossils indicating predation,
include two-dimensional, bedding-parallel, and include the holes drilled in the shells of other
three-dimensional forms. The key morphological molluscs by some gastropods by mechanical and/or
element is the derivation of a series of branching, chemical means (Figure 7M); the acid secreted by
non-interpenetrating, straight or curved shafts, or muricid gastropods can have a pH as low as 3.8.
spreiten-infilled lobes, from a common source (Figures More equivocal examples in soft substrates include
7F–7H). Domichnia are open burrow systems, or the intersection of two trace fossils, following which
borings into xylic and lithic substrates (Figures 7I– only one continues.
7L), utilized on a semipermanent basis; most are for This classification scheme works well for the ma-
habitation, but structures specifically constructed jority of trace fossils, although it has its limitations.
for other functions (e.g., brood chambers) have been Categories can grade into each other; for example,
526 TRACE FOSSILS

Figure 7 Variation in behaviour represented by trace fossils. (A) The cubichnion, Rusophycus. (B) An arthropod-produced repich-
nion. (C) The pascichnion, Nereites, exhibiting strophotactic and thigmotactic behaviour. (D) The consistent sense of braiding
between successive burrows indicates thigmotactic behaviour achieved by spiralling alone. (E) The agrichnion, Paleodictyon.
(F) Fodinichnion comprising unbranched shafts radiating parallel to bedding from a central area. (G) Fodinichnion comprising a
series of lobate spreiten. (H) High-density, monospecific occurrence of the fodinichnion, Chondrites; each burrow system comprises
a series of branching shafts (at arrows) fanning outwards and downwards from a central shaft. (I) Protrusive spreite (short arrows)
contained inside a U-shaped burrow (long arrows). (J) Oblique view of block of shallow marine oolitic limestone that was later
bored and encrusted during emplacement of a hardground community. (K) External view and rare example of a pupa preserved
inside the case constructed by a caddis fly larva. From Hugueney M, Tachet H, and Escuillié F Caddisfly Pupae from the Miocene
indusial Limestone of saint-Gerand-le-Puy, France (1990) Palaeontology 33: 495–502, Plate 1, Figures 3 and 4. (L) Bioglyph on the
internal surface of a burrow excavated in semilithified sediments. Image courtesy of Richard Bromley, Copenhagen. (M) Praedichnion:
a hole drilled by a gastropod through the shell of a bivalve. Scale bars, 5 mm.

at what point is a meandering pattern sufficiently Use as Environmental Indicators

regular to warrant the descriptor pascichnion, not
repichnion? Opinions may vary between authors Information from trace fossils and ichnofabrics can
as to what the dominant behaviour is; the burrow be incorporated into palaeoenvironmental recon-
Planolites has been considered a fodinichnion, pas- structions at scales ranging from the individual bed
cichnion, and repichnion, as it involves reworking of (reconstruction of a single endobenthic community)
the sediment whilst on the move in a straight to to depositional sequences tens to hundreds of metres
sinuous curve. Finally, each ichnospecies within an thick (e.g., ichnofacies).
ichnogenus need not have the same ethology; more
Ichnofacies
rarely, the ethology may vary (e.g., between a repich-
nion and a pascichnion) amongst a set of specimens of Ichnofacies are recurrent combinations of sediment-
an ichnospecies. ary facies and trace fossils; the Skolithos, Cruziana,
 TRACE FOSSILS 527

Zoophycos, and Nereites ichnofacies, emplaced than depth controlled. Environmental conditions are,
within marine softgrounds, characterize successively however, far from uniform at any given water depth.
greater water depths (Figures 9 and 10). The use of Local factors may control the distribution of ichno-
trace fossils as a palaeobathymetric indicator exploits facies; for example, depositional conditions in the
the fact that many, particularly ichnogenera, have a proximal or channellized parts of a submarine fan
long time range, but are restricted to, or most com- may resemble the high-energy, mobile substrates typ-
mon in, specific environmental conditions. These ical of nearshore environments, and the Skolithos
conditions include wave or current energy, tempera- ichnofacies occur in each. A change in environmental
ture, chemistry (including salinity and quantity of conditions can produce a succession of ichnofacies
dissolved gases such as oxygen), light penetration, that mimics, but is not the product of, significant
nutrient supply, competition for ecospace and re- changes in water depth; progradation of the sub-
sources, sedimentation rate, and substrate character marine fan in Figure 9A would result in lithologies
(including the grain size and geotechnical properties with a Skolithos ichnofacies succeeding those with a
of soft sediments). Changes in these conditions tend Nereites ichnofacies, unaccompanied by any signifi-
to correlate with changes in absolute water depth, cant decrease in water depth. The Glossifungites,
and thus the palaeobathymetry is depth related rather Trypanites, and Teredolites ichnofacies are emplaced
into firmgrounds, lithic substrates, and xylic sub-
strates, respectively (Figure 10); water depth is not a
controlling factor (Figure 9B). Some authors recog-
nize the Psilonichnus (between the foreshore zone and
the terrestrial realm) and Arenicolites (opportunistic
colonization of newly deposited event beds) ichnofa-
cies. Originally identified as the Scoyenia ichnofacies,
the heterogeneity of non-marine environments does
not lend itself to classification; several alternative
detailed subdivisions have been proposed, but no
consensus has emerged.
The presence or absence of an individual ich-
nogenus, even that after which the ichnofacies
Figure 8 Modification of the idealized vertical stratification is named, is not strong evidence for a particular
of electron acceptors via the emplacement of an open burrow water depth. Furthermore, the bathymetric ranges of
structure at depth. some ichnotaxa are known to have changed over

Figure 9 Schematic representation of the distribution of ichnofacies in marine environments. (A) Passive continental margin.
(B) Sediment-starved active continental margin. Reprinted from Bromley RG and Asgaard U (1991) Ichnofacies: a mixture of
taphofacies and biofacies. Lethaia 24: 153–163 (www.tandf.no/leth), by permission of Taylor and Francis AS.
528 TRACE FOSSILS

Figure 10 Summary of the ethologies, lithologies, and sedimentary processes characteristic of each of the main ichnofacies. After
Frey RW and Pemberton SG (1984) Trace fossil facies models. In: Walker RG (ed.) Facies Models, 2nd edn., pp. 189–207. Ontario:
Geological Association of Canada.
 TRACE FOSSILS 529

time. During most of the Palaeozoic, the ichnogenus is heterogeneous and occurs as discrete burrows.
Zoophycos occupied a broad range of marine water These are usually well defined because of the high
depths; since the Early Permian, it has become pro- shear strength of these more dewatered sediments
gressively restricted to greater water depths and is (softgrounds or even firmgrounds); other progressive
only found in continental slope and deep basin changes with depth include reductions in both the
settings today. volume and degree of oxygenation of the interstitial
pore waters. Vertical partitioning, tiering, of the tran-
Use of Infaunal Ecospace: Endobenthic Tiering
sition layer infauna (and thus the trace fossils they
Bioturbation in modern, fine-grained substrates emplace) occurs in response to such changes in the
undergoing accretion that is the work of a single physical and chemical properties of the sediments
community can be divided into three general levels: (Figure 11A). Reconstruction of the tiering profile
the surficial mixed and underlying transition layers therefore provides a measure of the community com-
in which bioturbation occurs, and the lowest histor- plexity; this can include the depth to which sediments
ical layer which contains the ichnofabric preserved were bioturbated and thus the volume of ecospace
after diagenesis and lithification (Figure 11A). As se- exploited. As sediment accretes, organisms will move
diment properties change progressively with depth, upwards to maintain the same level or ecological
the boundary between the mixed and transition layers niche; trace fossils produced at shallower depths will
is gradational, not abrupt; it is convenient, however, thus be cross-cut by those emplaced at successively
to model the two as distinct layers. Mixed layer sedi- greater depths. Dominant to unilateral cross-cutting
ments are often soupgrounds and thus, although of one trace fossil by another implies that the latter
bioturbated completely, the trace fossils are often belonged to a deeper tier; the mutual intersection
poorly preserved. Bioturbation in the transition layer of two or more trace fossils implies that they are

Figure 11 Endobenthic tiering revealed by the ichnofabrics produced 1. (A) Subdivision of the sediment column into the mixed (M),
transition (T), and historical (H) layers; inset shows dominant to unilateral cross-cutting relationships amongst the transition layer
trace fossils allowing the identification of four tiers (t1–4). (B, C) Rapid relocation of the infauna following event bed deposition leaves a
‘frozen profile’ in the older sediments, all of which (B) or, if the event bed is erosively based, only the lower part of which (C), may be
preserved. (D) Production of a ‘frozen profile’ resulting from the evacuation of the sediment column by infauna following a rapid
deoxygenation event (RDE). (E) A gradual deoxygenation event (GDE), in which the oxygen content of the interstitial pore waters
declines gradually, results in exclusion of the infauna of successively deeper tiers. Adapted from Savrda CE and Bottjer DJ (1986)
Trace-fossil model for reconstruction of paleo-oxygenation in bottom waters. Geology 14: 3–6.
530 TRACE FOSSILS

components of the same tier. In practice, deviations

from the random cross-cutting of older structures by
younger can occur; these include preferential re-
exploitation as well as avoidance of earlier formed
structures. Complete reworking of sediment at depth
will obliterate the record of earlier activity in shal-
lower tiers, and thus reduce the diversity of the trace
fossil community (the palaeoichnocoenosis). The pre-
servation of a complete tiering profile requires the
intensity of bioturbation to decline with depth; earlier
formed parts of the ichnofabric, including occasion-
ally the mixed layer ichnofabrics, occur as relict
patches between later structures.
The deposition of an event bed will result in a rapid
upward relocation of the infauna; if this event bed
is sufficiently thick, the base of the transition layer
will be moved above the older sediments, leaving
a frozen tiering profile preserved within them; the
distance from the sediment surface to the base of
the transition layer indicates the thickness of sedi-
ment occupied (Figure 11B). The upper part of the
sediment column is often a soupground, and thus
remobilized relatively easily; the upper parts of
the frozen tiering profile, notably the mixed layer,
may be eroded during the deposition of the next bed
(Figure 11C). Evacuation of the sediment column
following a rapid deoxygenation event, in which the Figure 12 Endobenthic tiering revealed by the ichnofabrics
redox threshold boundary is moved above the sedi- produced 2. (A) The base of the event bed cannot be reached
by the infauna of the shallower tiers; the trace fossils at its
ment–water interface, will leave a frozen profile pre- sole will comprise those of tier 3 and, if the accretion of sedi-
served below the succeeding sediments (Figure 11D). ment subsequently is gradual, tier 4. (B, C) Emplacement of
Mixed layer ichnofabrics, rare in the geological secondary trace fossils defines a piped zone (pz), the thick-
record, are more likely to be preserved in this manner ness of which is a measure of the volume of ecospace utilized.
than by burial below an event bed. For clarity, earlier formed parts of the ichnofabrics have been
omitted.
Other means of reconstructing the tiering profile
include the identification of the depth to which an
intergenic postdepositional trace fossil penetrated the
substrate. In a sequence of event beds of different and pelagic sequences are characterized by alternat-
thickness, the components of shallower tiers occur ing lithologies, the differences in colour and compos-
only on the soles of thinner beds (Figure 12A). Sec- ition of which can be the result of orbital forcing. In
ondary trace fossils (see above) occur within a ‘piped such cases, trace fossils within the piped zone may be
zone’ (Figures 12B and 12C). The thickness of the actively infilled by the same sediment as the host
piped zone indicates the volume of ecospace used by lithology.
the endobenthic community; components of progres- Natural systems are obviously more complex, but
sively deeper tiers occur closer to its base. In the more dynamic models can be produced by allowing
simple model in Figure 12B, the secondary trace variables, such as the rate of sediment accretion,
fossils were sourced from the immediately overlying including negative values (erosion), or the levels of
layer whilst it was being deposited, and the piped oxygenation, to fluctuate over time. Not all circum-
zone is contained entirely within one layer. The exam- stances will satisfy the assumptions within the model.
ple in Figure 12C shows a more complex situation; Thin-bedded siliciclastic turbidites often exhibit a
the lowest bed was rebioturbated during deposition bipartite division into a sand-rich lower part and a
of the second, not the first, overlying layer, but the mud-rich upper part, and infauna can position them-
latter was too thin to contain the piped zone entirely. selves with respect to the interface between the two,
In this idealized example, the colour of each layer is independent of its depth below the sediment–water
different, and the source of sediment in the piped zone interface. Cross-cutting relationships can also be
is obvious. In practice, however, many hemipelagic generated by the superimposition of two successive
 TRACE FOSSILS 531

Figure 13 Definitions and schematic illustrations of ichnofabric indices for strata deposited in: (A) shelf environments; (B) high-
energy nearshore sandy environments dominated by Skolithos; (C) high-energy nearshore sandy environments dominated by Ophio-
morpha; (D) deep-sea deposits, including examples of homogenized sediment that has been reworked subsequently. Reprinted with
permission from Droser ML and Bottjer DJ (1993) Trends and patterns of Phanerozoic ichnofacies. Annual Review of Earth and Planetary
Sciences 21: 205–225. ß 1993 by Annual Reviews www.annualreviews.org

communities; following event bed deposition, it is Intensity of Bioturbation

not unusual for a short-lived opportunistic com-
The ichnofabric index (ii) is a semiquantitative meas-
munity to exploit the new ecospace and resources,
ure of the intensity of bioturbation, based on the
before the longer term equilibrium community is extent to which the original stratification is disrupted;
re-established.
as both the nature of bioturbation and host lithology
Palaeo-oxygenation will vary, the schematic illustrations are specific to
certain environmental settings (Figure 13). The ich-
As bottom water oxygenation declines, the general
nofabric is observed in vertical sections; a standard
tendency is for the thickness of the mixed and transi-
horizontal field of view should be used and reported
tion layers, and the size (reflected in the burrow
(500 mm is often used in outcrop studies).
diameter) and diversity of actively infilled transition
layer trace fossils, to decrease (Figure 11E). It has
been suggested that ichnofaunal assemblages dom- See Also
inated by domichnia, pascichnia, and fodinichnia
Biosediments and Biofilms. Diagenesis, Overview.
indicate a progressive decline in pore water oxygen-
Fossil Invertebrates: Trilobites. Palaeoecology. Palae-
ation. Related to this, certain ichnogenera, notably ontology. Sedimentary Environments: Depositional
the fodinichnion Chondrites, especially if abundant Systems and Facies; Storms and Storm Deposits.
in a low-diversity or monospecific assemblage, have
been considered to be diagnostic of low oxygen con-
ditions; however, this should not be assumed without Further Reading
supporting ichnological and sedimentological evi- Bottjer DJ, Hagadorn JW, and Dornbos SQ (2000) The
dence. The shaft diameter is relatively large in the Cambrian Substrate Revolution. GSA Today 10: 1–7.
monospecific assemblage of Chondrites emplaced in Bromley RG (1996) Trace Fossils. Biology, Taphonomy and
a storm bed in Figure 7H, and an interpretation as Applications. London: Chapman and Hall.
an opportunistic colonization of newly deposited Bromley RG and Asgaard U (1991) Ichnofacies: a mixture
sediment is favoured. of taphofacies and biofacies. Lethaia 24: 153–163.
532 TRACE FOSSILS

Donovan SK (ed.) (1994) The Palaeobiology of Trace Maples CG and West RR (eds.) (1992) Trace Fossils. Short
Fossils. Chichester: Wiley. Courses in Paleontology No. 5. Tulsa, OK: Paleonto-
Droser ML and Bottjer DJ (1993) Trends and patterns of logical Society.
Phanerozoic ichnofacies. Annual Review of Earth and Pemberton SG (1992) (ed.) Applications of Ichnology to
Planetary Sciences 21: 205–225. Petroleum Exploration – A Core Workshop. SEPM
Ekdale AA, Bromley RG, Pemberton SG (1984) Ichnol- Core Workshops No. 17. Tulsa, OK: Society of Economic
ogy. The Use of Trace Fossils in Sedimentology and Paleontologists and Mineralogists.
Stratigraphy. SEPM Short Course No. 15. Tulsa, OK: Savrda CE (1995) Ichnologic applications in paleocea-
Society of Economic Paleontologists and Mineral- nographic, paleoclimatic and sea-level studies. Palaios
ogists. 10: 565–577.
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models. In: Walker RG (ed.) Facies Models, 2nd edn., reconstruction of paleo-oxygenation in bottom waters.
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