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Analysis and application of ichnofabrics

Article in Earth-Science Reviews · February 2003

DOI: 10.1016/S0012-8252(02)00105-8

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 Earth-Science Reviews 60 (2003) 227 – 259
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Analysis and application of ichnofabrics

Andrew Taylor a, Roland Goldring b,*, Stuart Gowland a
a
Ichron Ltd, 5 Dalby Court, Gadbrook Business Centre, Rudheath, Northwich, Cheshire CW9 7TN, UK
b
Postgraduate Research Institute for Sedimentology, University of Reading, Whiteknights, P.O. Box 227, Reading RG6 6AB, UK
Received 22 February 2002; accepted 4 June 2002

Abstract

Bioturbation at all scales, which tends to replace the primary fabric of a sediment by the ichnofabric (the overall fabric of a
sediment that has been bioturbated), is now recognised as playing a major role in facies interpretation. The manner in which the
substrate may be colonized, and the physical, chemical and ecological controls (grainsize, sedimentation rate, oxygenation,
nutrition, salinity, ethology, community structure and succession), together with the several ways in which the substrate is tiered
by bioturbators, are the factors and processes that determine the nature of the ichnofabric. Eleven main styles of substrate tiering
are described, ranging from single, pioneer colonization to complex tiering under equilibria, their modification under
environmental deterioration and amelioration, and diagenetic enhancement or obscuration. Ichnofabrics may be assessed by
four attributes: primary sedimentary factors, Bioturbation Index (BI), burrow size and frequency, and ichnological diversity.
Construction of tier and ichnofabric constituent diagrams aid visualization and comparison. The breaks or changes in
colonization and style of tiering at key stratal surfaces accentuate the surfaces, and many reflect a major environmental shift of
the trace-forming biota due to change in hydrodynamic regime (leading to non-deposition and/or erosion and/or lithification),
change in salinity regime, or subaerial exposure. The succession of gradational or abrupt changes in ichnofabric through
genetically related successions, together with changes in colonization and tiering across event beds, may also be interpreted in
terms of changes in environmental parameters. It is not the ichnotaxa per se that are important in discriminating between
ichnofabrics, but rather the environmental conditions that determine the overall style of colonization. Fabrics composed of
different ichnotaxa (and different taphonomies) but similar tier structure and ichnoguild may form in similar environments of
different age or different latitude. Appreciation of colonization and tiering styles places ancient ichnofabrics on a sound process-
related basis for environmental interpretation.
D 2002 Elsevier Science B.V. All rights reserved.

Keywords: ichnofabric; colonization; tiering; sequence stratigraphy; event beds; genetic successions

1. Introduction where primary sedimentary structures have been

The value of trace fossils in facies interpretation is
now widely accepted, particularly in successions
* Corresponding author.
E-mail addresses: taylor@ichron.com (A. Taylor),
r.goldring@reading.ac.uk (R. Goldring).
partly or wholly obscured by animal and/or plant
activity at all scales. Compared with body fossils,
their overall information potential is both more spe-
cific (due to the fact that trace fossils are rarely
reworked) and have other applications, for example,
in the investigation of porosity/permeability relation-
ships in highly bioturbated reservoir strata. The appli-

0012-8252/02/$ - see front matter D 2002 Elsevier Science B.V. All rights reserved.
PII: S 0 0 1 2 - 8 2 5 2 ( 0 2 ) 0 0 1 0 5 - 8
228 A. Taylor et al. / Earth-Science Reviews 60 (2003) 227–259
cation of trace fossils to facies interpretation has
developed since Seilacher (1954, 1955) appreciated
the major distinction between the ichnology of deep-
water turbiditic sands and muds, and marginal marine
facies. Where primary sedimentary structures have
been all but eliminated over an appreciable thickness,
an analysis of the ichnofabric (the overall texture of a
bioturbated sediment: Bromley and Ekdale, 1986;
Bromley, 1996) is the only practicable way to inves-
tigate the primary sedimentology and to obtain clues
about the depositional environment and its diagenetic
history. Ichnofabrics, by their rather inscrutable and
enigmatic nature are puzzling for geologists. They
neither readily relate to dynamic attributes nor resolve
into clearly identifiable and interpretable trace fossils.
The objectives of this review are to focus on the
factors and processes that determine ichnofabrics and
show how they can be interpreted and utilised in both
facies interpretation and stratigraphic correlation.
Some authors have used the term ichnofabric quite
loosely as an appendage to a simple listing of trace
fossils, but without any serious analysis of primary
sedimentology, or the spatial and temporal relation-
ships between the contained ichnotaxa. This approach
is to be regretted.
While ichnofacies are characterized essentially by
their assemblages of ichnotaxa (Bromley, 1996, p.
Table 1
The controls and attributes of ichnofabrics
Colonization and colonization strategy
Controls:
(i) Ethology and method of fabrication
(ii) Grain size
(iii) Nature of substrate
(iv) Oxygenation
(v) Nutrient levels
(vi) Salinity
(vii) Sedimentation rate
(viii) Colonization Window
Tiering pattern and ordering
Attributes:
(i) Primary sedimentological factors
(ii) Burrow size and frequency
(iii) Bioturbation Index
(iv) Ichnodiversity
Related aspects
Trace fossils and ichnocoenoses, ecological succession
Ichnoguilds
Compactional and diagenetic factors
290), ichnofabrics have been to a large extent distin-
guished by pattern recognition and position within
genetically related successions (Bromley, 1996, p.
294). Enhancement or diminution of the secondary
fabric may subsequently follow due to diagenetic
processes, and the latter are especially marked in
calcareous sediments. Of course, it is the ichnotaxa
themselves and their cross-cutting relationships (order-
ing) that most characterize the ichnofabrics, together
with distinctive morphologies, such as the pellet-lining
of Ophiomorpha (Fig. 8) or the frogspawn appearance
of Phycosiphon (Fig. 7). Although not excluded, shal-
low-penetrating epigenically formed taxa do not play
an appreciable role in the formation of ichnofabrics,
especially when viewed in core.
The element of the fabric of a sediment that is due
to bioturbation may be attributed to a number of
controls and attributes (Table 1, Fig. 1), in particular,
to the tiering pattern and manner of colonization.
2. Colonization and colonization strategy
The origin of the colonizing infauna is an aspect
that is not readily resolved: whether it arrived at a
colonization site as larvae or spat, or as ‘adults’ in the
process of relocation (Fig. 1). Ophiomorpha in deep-
marine sandy (and typically proximal) turbidites is
often regarded as indicative of relocation (Föllmi and
Grimm, 1990). Uchman (1995) regarded occurrences
in Miocene flysch of northern Italy as a normal
component of the deep-sea ichnoassemblage, since
burrow size distribution indicates that complete life
cycles are represented. Further, Uchman (1995) and
Wetzel and Uchman (2001) show that from a study of
cm-thick closely spaced turbidites, Ophiomorpha was
formed by producers moving upwards. It is also
recognised that very few animals occupy a single
life-long position within the substrate (Schäfer,
1972; Frey and Goldring, 1992, Fig. 3). What is
important, however, are changes in ichnofabric that
occur gradually through a succession, or sharply at a
key stratal surface, or take place across an event bed.
These are discussed below.
Animals and plants do not create uniform distribu-
tions of burrows, tunnels or roots, but a patchiness
that can to some extent be evaluated in ancient
successions, when outcrop allows. Uchman (2001)
 A. Taylor et al. / Earth-Science Reviews 60 (2003) 227–259 229

Fig. 1. General figure to show collectively the various controls (left) and attributes (right) of an ichnofabric, from primary sedimentation to
diagenesis. Controls: Manner of colonization: larval settlement or relocation. Origin of biota: (A) from substrate surface; (B) from an eroded
(and thus firmer) surface; (C) from below, by upward migration; (D) from a level substantially above, through sediment subsequently
penecontemporaneously eroded. Ethology, nutrition and oxygenation of the substrate at different depths below the substrate surface (see Figs. 3
and 4). State of substrate (see Fig. 2), sedimentation rate. Colonization Window (see Section 3.8). Tiering and ordering (see Fig. 6), here
illustrated by a frozen tier associated with tiered ichnotaxa. Diagenesis, involving compaction, enhancement (as seen with Ophiomorpha
galleries enhanced by preferential lithification, while unlined shafts remain less distinct) or obscuration of burrows within an early cemented
concretion. Attributes: Bioturbation Index (BI) (see Table 3). Ichnofabric Constituent Diagram (see Section 5.2 and Fig. 7). Burrow size (most
conveniently given as burrow diameter), and burrow frequency (see Section 5.3). Diversity of ichnotaxa (see Section 5.4). Tier diagrams (as
cartoon) to indicate the ichnotaxa, and relative level at which they produced an ichnofabric (see text and Table 4).

described the patchiness of trace fossils in flysch hardground. But, more often, when dealing with core,
deposits, including basin-plain environments. Also, one trusts that over the time interval represented by
Palmer and Palmer (1977) were able to evaluate the core, a ‘time-averaging’ of the traces will have
patchiness on an encrusted and bored Ordovician been achieved.
230 A. Taylor et al. / Earth-Science Reviews 60 (2003) 227–259
Any burrow we see may have been initiated at one
of four possible sites (Fig. 1):
(1) At a substrate surface that can be positively
identified, e.g. funnel opening, or a complete bivalve
crypt (Fig. 1A).
(2) At a surface that had been penecontempora-
neously eroded and/or lithified prior to colonization.
The question will then be: can the amount of erosion
and degree of lithification be determined or estimated
(Fig. 1B)?
(3) The trace may be due to an organism entering
the sedimentary layer by moving upwards (Fig. 1C).
Then the question is: was the producer escaping, or
adjusting its position in response to gradual accretion,
or simply a predator moving through the sediment?
(4) The trace may be the result of an animal
burrowing down deeply from a higher level (probably
subsequently eroded) and into sediment that was
suitable, but which had been deposited under quite
different environmental conditions (Fig. 1D). This is a
common situation leading to deep burrows of Ophio-
morpha (Pollard et al., 1993) which may extend to a
metre and more below the substrate surface.
Under more or less continuous sedimentation and
colonization the manner of colonization, the method of
sediment penetration (intrusion, compression, excava-
tion: Bromley, 1996) can often be determined (Frey and
Goldring, 1992) from bioglyphs or by the style of
deformation of the host sediment. What is important
for facies interpretation is appreciating whether or not
the surface colonized genetically relates to the host
sediment. Frey and Goldring (1992) attempted to
resolve their problem by recording the colonization
surface in sedimentary logging rather than the biotur-
bated interval. In practice however, misinterpretation
only arises with deeply formed burrows or where there
has been significant penecontemporaneous erosion.
There are four strategies of colonization: the first
being the opportunistic, rapid colonisation of a vacant
ecological niche, e.g. following an overbank spill to
the floodplain, or deposition of a storm event bed or
tephra (volcanic ash) layer. Opportunists are of low
diversity and tend to be of small size, and hence
generally display small diameter burrows. Their life
span also tends to be short, so that complex (often
large) burrow systems are not formed. Providing
another event does not return conditions to ‘square
one’, the ‘generalist’ pioneers will be followed by a
more diverse biota leading to an equilibrium situation
and then to a mature climax community. The latter
may include complex burrow systems, e.g. Ophio-
morpha boxworks, and Zoophycos.
3. Controls on colonization
3.1. Ethology and method of trace fossil fabrication
Understanding how trace fossils are initiated and
their subsequent expansion and extension on and
within the sediment (the method of fabrication), and
for what purpose (ethology), are crucial for any real
appreciation of the role of a trace and, of course, to its
taxonomy. The ethological classification of trace fos-
sils developed by Seilacher (1954, 1955) is well-
known. Subsequent additions are documented in
Bromley (1996). It was only by careful observations
on Limulus by Caster (1938) that showed that the
culprits of Devonian trackways were not vertebrates
but more probably a xiphosurid. Analysis of much
material of Cruziana and allied traces by Seilacher
(1955) showed how the animal responsible, in most
cases a trilobite, moved over the substrate surface in
various ways, sometimes being deflected by currents,
or systematically digging into the substrate to produce
the various traces that can be observed and utilised
stratigraphically (Seilacher, 1970, 1991, 1994). More
recently, Seilacher and Seilacher, 1994 and Seilacher-
Drexler and Seilacher (1999) made elegant experi-
ments on modern gastropods and other animals in
order to understand what happens within the sediment
(including the making of undertraces), especially at
substrate boundaries (bed junction preservation).
Behaviour and substrate consistency (below) are
inextricably linked, whether in respect of movement
and penetration, or feeding. While the substrate is
always affected by animal activity, the actual activity
is not necessarily controlled by substrate state, as with
large vertebrates moving over a substrate of somewhat
varied cohesiveness. But at a smaller scale, the state of
the substrate much determines the type of activity
possible, for example, whether or not the Ophiomor-
pha-producer ramifies its burrow margin.
An aspect of ethology that has scarcely been
addressed is ‘anomalous behaviour’. For example,
the basal spreading of Diplocraterion at a concealed
 A. Taylor et al. / Earth-Science Reviews 60 (2003) 227–259
firmground junction. Under experimental conditions
simulating bed-junction preservation, Jensen and
Atkinson (2001) noted that crustaceans burrowing
through a thin sand were perturbed on encountering
the underlying relatively firm mud (in fact, an artificial
concealed bed junction), by meandering along the
interface in a manner not previously observed in the
laboratory. This raises the spectre that most, or many
interface trace fossils (on which ichnology so much
depends) represent anomalous behaviour! Jensen and
Atkinson (op. cit.) referred to such traces as ‘barrier
prospecting traces’. Burrowing through the interface
would generally negate such interpretation. Graded
muds, with generally a thin cross-laminated basal layer,
are a common feature of lower shoreface-transitional
zone facies (Goldring, 1971; Walker and Plint, 1992)
associated with storm event beds. Only exceptionally
were such beds colonized, and associated load-cast
ripples testify as to the softness of the substrate.
Although a concealed bed-junction generally
means a grain-size change, geochemical changes, as
at the redox boundary, also represent ‘concealed’
junctions. ‘Normal’ behaviour is probably best repre-
sented where conditions favoured successive tiering,
and where minimal soft sediment deformation has
taken place at lithological boundaries. Miller and
Vokes (1998a,b) usefully distinguish between traces
formed incidentally and only occupied in the short-
term, from deliberate traces, representing relatively
long-term occupation and necessitating burrow mod-
ification, both during occupation (as with recycling of
the burrow contents), and modifications due to exter-
nal factors (for example, repair of openings, construc-
tion of multiple openings, or dealing with sediment
influx). Skolithos may be cited as an example of the
first, and Zoophycos and Phymatoderma examples of
the second. Nevertheless, interpretation of certain
ichnotaxa, notably Zoophycos, remains a much-
debated enigma (Gaillard et al., 1999; Bromley,
1991; Lockair and Savrda, 1998a,b). Fürsich (1973a,
1974a,b), defended by Schlirf (2000), placed various
ichnotaxa into synomymy, arguing that they displayed
the same basic fabrication. However, Seilacher (1967,
and pers. comm., 2000) has argued that although this
might be true, in the case of the long Rhizocorallium
irregulare related to sediment-ingesting activity, and
the short and often oblique Glossifungites jenense (a
trace fossil probably used for dwelling and filter
231
feeding), the differences are such as to warrant sepa-
ration at ichnogeneric level. This is not the place to
discuss ichnotaxonomy, but it is useful to note that
there is no formalised method of naming ichnofabrics.
Ethology and the ethological spectrum of sedimen-
tary facies were major aspects for Seilacher (1954,
1955) in distinguishing between deep-sea and shal-
low-marine traces. Differences in the ethological
spectra are emphasised by the low preservation poten-
tial of surface feeding traces in shallow water sedi-
ments. The problem in slabbed core is aggravated
further by the lack of evidence for bedding parallel
burrows, especially interface traces, that are generally
prominent at outcrop, especially in heterolithic lithi-
fied sediments of many facies. Further, interpretation
of the trophic group to which traces such as Skolithos
and Diplocraterion belong can be uncertain: whether
the burrows represent the domicile of a suspension or
deposit feeder. Documentation of stratigraphical
changes in the ethological and trophic group spectra
(at all scales), however, is useful as supporting evi-
dence in environmental interpretation.
3.2. Grain size
Larval settlement (Butman, 1987) and the manner in
which animals penetrate the sediment is obviously
much influenced by grain size and packing, as is the
way animals feed on or manipulate grains (e.g. Warren
and Sheldon, 1968; Goldring, 1995). For example,
some crustaceans are restricted to medium-grained
sand because they have evolved the means to handle
and clean grains of this size. But it is now clear that in
respect of larval settlement, it is the hydraulic regime
rather than the substrate factor per se that is important.
In sandy substrates vertical shafts are typical, many
representing sites of likely suspension feeders.
Grain size is generally readily estimated in the
field, but a grain-size distribution derived from highly
bioturbated sediment can have no hydraulic signifi-
cance. Likewise, bioturbation, in destroying primary
sedimentary structures and the primary fabric, also
destroys the directional permeability associated with
these. Strongly bioturbated intervals will often have
reduced permeability, but the mixing-in of a propor-
tion of mud seems often to have hindered cementa-
tion. However, if ‘fines’ are expelled at the water
column by deposit feeders, e.g. the bivalve Yoldia
232 A. Taylor et al. / Earth-Science Reviews 60 (2003) 227–259

(Rhoads, 1963), rather than being pushed aside, then
the sediment will be cleaned. In partly bioturbated
sediments primary grain-size distribution can be
measured if care is taken to sample only areas
exhibiting primary sedimentary structures. (In lithified
rock this can be done on a thin-section: 100 grain
measurements may be sufficient.)
3.3. Nature of the substrate
Trace fossils and bioturbation provide the clearest
indications for the state of the substrate (soft, loose,
cohesive, stiff, firm, hard) at the time of their con-
struction (Fig. 2) (Goldring, 1995), and are particu-
larly important in the recognition of omission and key
stratal surfaces. This also applies to any sequential
changes to the substrate associated with lithification
that might have taken place. A general indication of
the response by organisms to substrate (six main
states) and their identification is shown in Table 2.
The manner in which the substrate was penetrated
by burrowers and borers has been much discussed (see
Bromley, 1996). Soft substrates and hardgrounds can
nearly always be readily discriminated. Hard substrates
are best distinguished by their borings and nearly
always indicate omission; most borings indicating
feeding from above the substrate. Likewise soft-
grounds and loosegrounds are marked by the nature
of burrow margins and compactional features (Gold-
ring, 1995, 1999a). Stiffgrounds (Wetzel and Uchman,
1998b) refer to unlined tunnels in mudrocks which
have been passively infilled, and firmground burrows
have sharp margins and the presence of bioglyphs
(scratches, etc., on burrow margin). But different types
of animal cope with firm substrates by using different
means of excavation; compare the mechanical excava-
tion by an arthropod with the grinding action of a
bivalve. Ruffell and Wach (1998) did not consider the
biological processes involved in the formation of
Aptian firmgrounds and their examples of ‘mature’
and ‘immature’ firmgrounds refer to typical soft- and
loose-ground situations at simple omission surfaces.
Macroscopic and microscopic borings can provide
an indication of water depth if they can be attributed
to the action of specific borers, particularly those
associated with photosynthesising algoids (Bromley
and Asgaard, 1993a,b; Bromley, 1994; Perry, 1996,
1998a,b; Perry and Bertling, 2000). Thus, Gnathich-
nus, the work of algal-grazing echinoids, and Entobia
gonoides, the work of the sponge Cliona virides (with
algal symbionts) are restricted to depths of a few
metres. In the Neogene and more recent sediments,
if attribution can be made to extant taxa, then water
depths can be inferred, as for example with Gastro-
chaenolites formed by Lithophaga. Turbidity (Brom-
ley, 1994) and turbulence also play a role as, for
example, even very low levels of sedimentation can
inhibit or destroy boring sponges.

Fig. 2. Diagram to show types of biogenic response to loose (soft), firm and hard (lithified) substrates: (approx.  0.5 unless otherwise
indicated): (A) Nucula; (B) Chondrites; (C) Ophiomorpha; (D) Echinocardium; (E) Mercenaria/Arctica (  0.25); (F) Thalassinoides; (G)
Spongeliomorpha (with scratched burrow margin); (H) Petricola; (I) Lithophaga (insoluble grain on left side) (  1.0); (J) crinoid holdfast; (K)
oyster: (L) Trypanites; (M) encrusting bryozoans and serpulids on cemented burrow wall.
 A. Taylor et al. / Earth-Science Reviews 60 (2003) 227–259 233

Table 2
Substrate type: observations and identification
Observations Deductions Type of ground
In mudrocks and micrites, sediment highly diffuse, Sediment was more or Soupground
possible traces highly compressed and smeared, less water-saturated,
feeding burrows dominant, e.g. Planolites compaction extreme

In muddy and micritic sediment, burrows show Some dewatering Softground

substantial compaction and indefinite margins, or a
thin mud lining. Bioglyphs indistinct or not preserved.
Mixed feeding (in deepwater: farming traces),
Chondrites, Zoophycos, Phycosiphon, Helminthopsis

In sandy sediment, burrows lined: Ophiomorpha. Permanent burrows, requiring Looseground

Bivalve burrows, Arenicolites stabilised margins

In muddy sediment, unlined burrows stable e.g. Stabilised sediment Stiffgrounda

Thalassinoides

Burrow outlines sharp, and may have distinct sharp Stiff, dewatered substrate Firmgroundb,
bioglyphs (scratches). F slight deformation of crypts Concealed firmgroundc
and compaction of passively filled burrows:
Spongeliomorpha, Glyphichnus.
Some encrusters e.g. Liostrea

Mineralised crust ( F ). Boring and encrusting biota. Lithified substrate surface Hardground
Borers cut evenly across matrix and skeletal grains (in limestones),
Shellground
(cemented shell bed),
Rockground
(with tectonic omission)
a
Proposed by Wetzel and Uchman (1998b) as a category distinct from firmground which they restricted to mechanically excavated burrows
with organised bioglyphs.
b
This category is not well-defined.
c
Sharp change in behaviour, generally associated with lithological change indicating firm interface within the transition zone. The effect is
to tend to lead burrowing along the interface (barrier prospecting traces).

3.4. Oxygenation
Highly bioturbated sediment can represent no other
than a well-oxygenated sea floor (Figs. 2 and 3). If
finely laminated black mudrock represents the oppo-
site, then successions which show gradations one into
the other (Fig. 3), represent deteriorating or amelio-
rating oxygen gradients) (reviewed by Wignall, 1994;
Allison et al., 1995). Stratigraphic changes in oxy-
genation are readily recognised in the field, and
especially in slabbed core. Changes seen in the rock
have been shown to be close, if not identical, to
changes observed in modern deep-sea cores. They
chiefly concern, in respect to decreasing oxygenation:
(a) decrease in trace size (burrow diameter); (b)
decrease in trace diversity; (c) decrease in the number
of tier levels (below), with loss of higher tiers, and an
associated upward shift of tiers.
If Chondrites is present, all three criteria may be
met, but it is safer not to rely on a single taxon. Shift
of tier depth is obviously less well appreciated since it
will be related to sedimentation rate. Gradual change
probably took place over a considerable period of
time, in contrast with an oxygenation ‘event’ or
seasonal change. Slow re-oxygenation may favour
the formation of stabilizing microbial mats. Abrupt
re-oxygenation is probably due to bottom current
activity associated with (slight) erosion. Decrease in
burrow size with depth only relates to functionally and
behaviourally similar burrows. Besides oxygenation,
three other ecological factors should also be kept in
mind: the nature of the substrate, sedimentation rate
234 A. Taylor et al. / Earth-Science Reviews 60 (2003) 227–259

Fig. 3. (A,B) Deoxygenation: (A) Changes in tier structure with gradual deoxygenation (from observations on North Sea core); (B) a rapid
deoxygenation event (modelled on Wignall, 1994); (C,D) distinguishing between soft substrate and low oxygenation (after Wignall, 1994); (C)
faunal changes under a declining oxygen gradient and (D) in a fine-grained clastic substrate on transition from firm to soft conditions. Epibiont
species richness refers to hard, shelly substrates. Epifaunal taxa, left to right: Chlamys, Gryphaea with abundant epibionts including serpulids,
Parainoceramus, small Gryphaea, procerithid gastropod, Parainoceramus, serpulid, trochid gastropod, Gryphaea, ‘paper pecten’, Gryphaea.
Infaunal taxa, left to right: mecochirid crustacean, Goniomya, Nicanella, Pinna, Nucinella, Dentalium, Palaeonucula. There is no zero point in a
purely muddy situation.

and food supply, especially the availability of sus-
pended food.
3.5. Food availability
Food availability is not an important factor in
determining the distribution of trace-forming organ-
isms in shallow marine environments, though it might
be predicted that oligotrophic sites might show low
levels of bioturbation. But it is now recognised that
organic matter is not uniformly available in the deep
sea. Where oxygen deficiency is not a limiting factor
(Wetzel and Uchman, 1998a) high input of organic
matter is associated with a dark colour and relatively
deep tiering with diverse traces (Fig. 4). Brown to
green colour is associated with low sedimentation
rates. The evidence is especially clear in the frozen
tiers at the base of turbidites where minimal erosion
has occurred. As Wetzel and Uchman (1998a,b) dis-
cuss, if benthic food content is high, specialised
search behaviour is not required: traces tend to be
large and unguided, and extend deeply into the sub-
strate. Specialised behaviour is mostly realised by
shallow tier or epigenic (sediment surface) trace
makers, and organisms forming graphoglyptids (Pale-
eodictyon) (Fig. 4) under low organic matter input. It
 A. Taylor et al. / Earth-Science Reviews 60 (2003) 227–259 235

Fig. 4. Hemipelagic sedimentation where bottom-water oxygenation is not restricted, but with fluctuations in organic matter input, and in
sedimentation rate. Sediment colour indicated. (a) Low organic matter input and low sedimentation rate; thin bioturbated zone with small
diameter burrows and graphoglyptids common. (b) High organic matter input associated with low sedimentation rate, high densities of surface
grazing traces, more deeply penetrating burrows of small size and depth of bioturbated zone restricted. (c) Low organic matter input but with
high sedimentation rate, graphoglyptids common and bioturbated zone extended in depth, but burrows not size-reduced and burrow reworking
common. (d) High organic matter input and high sedimentation rate, with bioturbated zone extended downwards, burrows not reduced in size,
burrow reworking common, high densities of surface grazing traces, and overall diversity high (after Wetzel and Uchman, 1998a).

is the interpretation of the function of Zoophycos and,
to a lesser extent, Chondrites that remain the problem
for ichnology in general.
3.6. Salinity
The ichnology of brackish water facies is some-
what controversial and is discussed further. At present
we do not understand sufficiently the biological con-
trols, particularly how marine animals cope with
reduced salinity, both within species and between
species and groups. In tidal environments deep bur-
rowers find temporary refuge and can penetrate fur-
ther upstream than epifaunal animals. It is clear that
some animals (bivalves, e.g Mytilus in the Baltic)
become stunted in reduced salinities, but this does not
apply to annelids. Some arthropods indeed are able to
adapt physiologically without any decrease in size.
But since we can rarely identify the culprit of a trace
in ancient sediments we have no means of telling
whether its constructor could adapt. Organism size
(and hence burrow diameter) depends on many fac-
tors, including age and food availability, and occupa-
tion time.
236 A. Taylor et al. / Earth-Science Reviews 60 (2003) 227–259
A number of criteria have been taken to indicate
toleration to brackishness in ichnotaxa. There is no
disputing that there is a general reduction in diversity
from open marine conditions towards brackish con-
ditions. But the latter are generally associated with
high population densities; a feature of opportunistic
colonization. The food resources of estuaries and
lagoons are generally high. There are no ichnotaxa
that are specific to brackish water facies. But if an
integrated approach is adopted (ichnology, sedimen-
tology, palynofacies, etc.), including an analysis of the
Colonization Window (Section 3.8) then a useful
environmental interpretation can be made.
In lagoonal sediments, where a shelly macrofauna
may often be present, ‘salinity events’ may be indi-
cated by stratally contained occurrences of Diplocra-
terion (often indicating marine flooding surfaces in
Mesozoic lagoonal deposits; e.g. Goldring et al.,
1998; Taylor and Gawthorpe, 1993). In estuaries a
salinity gradient ranging from normal marine salinity
to fresh water is to be expected, and any sound
environmental interpretation requires a thoroughly
integrated approach.
3.7. Sedimentation rate
Sedimentation rate and the role of penecontem-
poraneous erosion have long been recognised as
major influences on bioturbation (Goldring, 1964;
Howard, 1975, whose models were drawn from mar-
ginal and shelf depositional environments). In deeper
water, sedimentation rate integrates closely with oxy-
genation and food availability. Many organisms can
adjust to an increase in the rate of sedimentation, but
to different degrees. The animal responsible for pro-
ducing Macaronichnus in the foresets of estuarine/
marine sand waves was particularly successful in
dealing with high rates of sedimentation. The main
effect of slow sedimentation on the ichnofabrics of
level bottom communities is in the overprinting of
burrows and borings. This renders interpretation of the
ordering and tiering difficult, and the burrows of
larger and deeper organisms (key bioturbators) are
over-emphasised. While event beds are ubiquitous,
more or less steady sedimentation of a high rate
appears to be, on a geological scale, less widespread,
or interbedded with event bed sedimentation. On a
level substrate the Cylindrichnus-producer (Goldring,
1996), with its exceptionally long apertural neck,
seems to have been particularly successful from the
Cretaceous onwards in conditions of high sedimenta-
tion rates, while Thalassinoides and allied taxa are
associated with apparently less rapid rates of sedi-
mentation.
3.8. Colonization Window
The duration of time potentially available for
colonization to take place, and for a burrow system
to be completed, the Colonization Window (Pollard et
al., 1993) is a significant factor. However, Pollard et
al. (op. cit.) were emphasising the physical processes
involved, and recognised the relatively short duration
of the Colonization Window under tidal conditions as
compared to an open marine situation. The often
incomplete morphologies of Ophiomorpha and low
Bioturbation Index (Section 5.2) in estuarine/delta-
distributary facies shows the Colonization Window to
be a major factor in the interpretation of this facies.
The duration of the Colonization Window reflects
environmental stability. Chemical aspects (Chemical
Windows) must also be a factor: salinity fluctuations
in the marginal marine environments and oxygenation
and fluctuations in food availability in lagoonal and
offshore environments (Wetzel and Uchman, 1998a).
These are less readily identified visually, but it is the
body and trace fossils that are likely to provide the
initial clues. Whilst in a marginal marine environment,
the duration of a Colonization Window can often be
determined with some confidence, the duration of
‘chemical windows’ is less certain.
3.9. Discussion
It is clear that the several controls that determine
the nature of the ichnofabric interact (one with the
other). Determining the relative importance of each in
ancient sediments is fraught with difficulty—as is well
appreciated in facies interpretation in general. Some
factors such as oxygenation have probably been over-
emphasised by workers, while Wetzel and Uchman
(1998a,b) showed that benthic food content has been
somewhat neglected. Temperature is another over-
looked ecological factor that has not seriously been
considered in ichnology in general. This control is
discussed further below. Further, the rate of change in
 A. Taylor et al. / Earth-Science Reviews 60 (2003) 227–259 237

the various controls is difficult to quantify or even
qualify in ancient sediments.
4. Tiering pattern and ordering
Partitioning of the substrate has long been appre-
ciated by biologists who refer to ecological stratifica-
tion below (or above) the substrate surface (or ground
level). Below the surface (Fig. 5), tier partitioning is
the result of animals (and plants) penetrating to
successive depths because of size and/or their needs
to utilize different feeding and respiratory strategies.
Bromley (1996) has reviewed modern studies on
muddy substrates demonstrating this phenomenon,
and made a distinction between the thoroughly bio-
turbated mixed layer, the transition layer (the zone of
maximum colour contrast with principal bioturbators),
and the underlying historical layer (without active
bioturbation). In thickness the mixed layer is generally
quite thin, and in muddy sediment may be no more
than a few millimetres. In contrast, the transition layer
can be several metres in thickness in inter-tidal/upper
shoreface sands. The mixed layer does not in effect
exist in mobile sandy substrates. The consistency of
the sediment, even within the transition layer, shows
changes in physical properties to which the burrowing
organisms may or may not be able to respond. This is
where concealed firmgrounds are particularly impor-
tant.
In viewing ancient sediments, the infaunal tier
patterns have been complicated by biological and
geological processes. In the former colonization may
have been patchy and/or fluctuated over time, raising
problems similar to those associated with the evalua-
tion of time-averaging shell beds (Kidwell and
Bosence, 1991). It is the processes of aggradation,
penecontemporaneous erosion, and accompanying
and succeeding geochemical and taphonomic pro-
cesses (lithification, etc.), and longer-term erosion
(unconformity), that chiefly affect the formation of
the ichnofabric. Gradual lithification may lead to

Fig. 5. Left, a modern tiered community in sand. A very shallow mixed tier is formed by small ‘worms’ and microorganisms, and a shallow-
burrowing starfish. The transition layer is occupied by bivalves and carnivorous gastropods, with an infaunal echinoid forming the deepest tier
at about 10 – 15 cm. The historical layer below is not colonised and sub-fossil. Based on Thorson (1968) and Bromley (1996). Right, the same
community as it might be, after slight erosion has removed the mixed layer and deposition of an event bed to form a frozen tier. With good
chance the molluscs might be preserved autochthonously. The Tellina is shown to have been drilled. Only the echinoid has left a distinctive
meniscate (orange-peel) backfill, and possibly a largely decorticated test. (N.B. Gastropod activity is more clearly preserved at a bed junction,
e.g. sand on mud).
238 A. Taylor et al. / Earth-Science Reviews 60 (2003) 227–259
successive replacement, as with hardground develop-
ment. Wetzel and Uchman (2001) showed that the
initial colonizers (Phycosiphon producer) of a muddy
turbidite utilizing a well-oxygenated substrate, were
replaced as the redox boundary became re-established
(closing the Colonization Window), by burrowers
utilising microbial matter just above the boundary
(Nereites producer), and the poorly oxygenated sub-
strate below (Chondrites producer). Thus, no cross-
cutting by Phycosiphon was observed. At the same
time deep burrowers unaffected by the event were
able to move upwards and to ‘reconnect’ with a new
substrate surface. Computer models of tiering and
bioturbation have been developed (Bromley, 1996),
which help to appreciate the complexity of the bio-
logical and geological processes. The effect of pene-
contemporaneous erosion in revealing tier structure
(as well as the amount of erosion) is well-known and
appreciated in shallow- and deep-water facies (e.g.
Wetzel and Aigner, 1986). In addition, evolutionary
changes, such as the colonization of deep-sea sedi-
ments by infaunal echinoids, and the appearance of
deep-burrowing strategies in the Bivalvia in the Mes-
ozoic have left distinctive effects on the sediment
fabric. In both, the impact of such key bioturbators
(animal species that by their activity dominate sub-
strate reworking) is to obscure other traces and reduce
the apparent diversity. The tier position of a particular
ichnotaxon can change with time and, probably,
between facies: a factor that must also be taken into
account. This is to be expected when it is appreciated
that identical types of trace fossils can be formed by
quite different organisms. Thus, there is an ongoing
debate on the role and tier position of Chondrites in
weakly oxygenated sediments (see review by Wignall,
1994).
The significance of tiering in ancient sediments is
threefold: for palaeobiology as a precursor to the
understanding of all processes affecting the deposi-
tional environment; in understanding the basis for the
observed porosity and permeability and, for the
understanding of subsequent consolidation and dia-
genesis.
Determination of the tier pattern in ancient sedi-
ments has generally been by three methods: (a) by
analogy with modern examples; (b) from observations
made below event beds, when minimal erosion has
taken place; (c) by cross-cutting relationships ob-
served in sediments deposited under more or less
continuous deposition (e.g. Frey and Bromley, 1984).
However, Lockair and Savrda (1998a,b) showed
that method (c) exaggerates the number of tiers. In a
study of limestone marl cycles they demonstrated that
a better estimate of the tier structure can be achieved
from observations of the maximum penetration depth
at lithological change where the overlying lithology
has been piped downwards. Thus, the seven tiers of
Frey and Bromley (1984) are better appreciated as
only two tiers. The effects of phobotaxis (behavioural
avoidance of contemporary burrows) and preferential
sediment exploitation are probably the main reasons
why direct measurement of activity depth provides a
better evaluation of the tiering. Depth per se is not the
dominant factor. Ordering, the relative timing of each
ichnotaxon’s introduction, can only be determined by
cross-cutting relationships.
Six styles of tiering were recognised by Goldring
(1995, 1999a). These are extended to eleven styles
(Fig. 6), where bioturbation is not significantly
affected by omission. Fig. 6a – e represent single
colonization events, though the burrowing may be
pervasive.
Single tier colonization by an opportunist or pio-
neer (Fig. 6a,b). Two types are shown. In (a), Skoli-
thos is illustrated as a deeply penetrating trace
generally associated with a sandy substrate. Vossler
and Pemberton (1988) reviewed the occurrence of
Skolithos as the trace of an opportunist. In (b) thin
beds (generally 20 mm or less in thickness) may be
completely riddled by the relatively shallow burrows
of Planolites in, for instance, lagoonal and flood-plain
facies where temporary environmental shifts have
allowed colonization (e.g. Pollard, 1988). Typically,
the mud-filled burrows represent the passage of the
burrower through a sandy unit from an overlying
mud.
Successive tiering by a single taxon in an aggrad-
ing depositional environment (Fig. 6c,d). In (c), the
successive tiering is exemplified by a bivalve, and is
shown by stacking (adjusting) under equilibrium con-
ditions. There is often an upward decline in the
number of individuals which may be attributed to an
environmental factor leading to death (Broadhurst et
al., 1980). The type of tiering is typical of a relatively
stable environment, but the low diversity suggests
stressed conditions. The ichnofabric is particularly
 A. Taylor et al. / Earth-Science Reviews 60 (2003) 227–259 239

Fig. 6. Eleven styles of tiering. (a – d) Four styles of colonization by a single taxon are shown: (a) by a shaft-producer; (b) by a deposit feeder
ramifying at a shallow depth; (c,d) aggradation by a continuously equilibrating mollusc, or a deep-tier arthropod periodically re-establishing
contact with the substrate surface. (e,f) Two styles of multiple colonization: simple tier with a single colonization, and complex tiering, where
several taxa create a fabric that is steadily overprinted as aggradation proceeds more or less continuously. Interruption may take place by erosion
or event bed sedimentation (Fig. 10) in a genetically related succession. (g – i) To illustrate tier modification with (g) tier replacement, (h,i)
tiering during amelioration (increasing O2) or deterioration (decreasing O2) (see also Fig. 3). (j,k) To illustrate enhanced and obscured fabrics by
diagenetic processes (see Fig. 1).

distinctive, and is characteristic of certain depositional
environments such as ‘sheltered’ sandy delta top
facies (e.g. crevasse splay). It is seldom evident in
shoreface or offshore facies because sedimentation
there tends to occur as pulsed events. Repeated levels
may be due to seasonal floodings (Eagar et al., 1985;
Hardy and Broadhurst, 1978). The colonization of
dune foresets in a marine facies, or on point-bar
surfaces in estuarine (or equivalent) facies, leading
to Macaronichnus or Palaeophycus, are variants in a
situation where a high sedimentation rate is readily
recognised. At the bases of foresets or between dunes,
where the sedimentation rate is lower and the sedi-
ment muddier, Macaronichnus may be overprinted by
other taxa.
Aggradation may not necessarily take place uni-
formly and continuously, but the evidence is shown by
buried stacked openings (cf. Frey et al., 1978) and has
been described from foreshore – shoreface environ-
ments (Fig. 6d). Adjustment downwards with relatively
slow but continuous erosion has also been described
from neritic facies (Bromley and Hanken, 1991).
Simple tiering by opportunists (or pioneers), rep-
resents a ‘one-off’ colonization (Fig. 6e). There is
likely to be minimal, or unrecognisable cross-cutting
by the ichnotaxa since only a single colonization
event is represented.
In these five situations, colonization appears as a
single event in a depositional environment, followed
by a unit of sedimentation (albeit only briefly).
240 A. Taylor et al. / Earth-Science Reviews 60 (2003) 227–259
Complex tiering can be attributed to aggradation
with more or less continuous colonization by the same
suite of taxa (Fig. 6f). Local differences in frequency
may cause changes in the vertical profile as viewed in
a core because of lateral restriction or patchiness of
colonization. If an event bed, without significant
erosion, has interrupted sedimentation, then a ‘frozen’
tier suite (Fig. 1) may be evident below the sole (i.e.
base), providing evidence of the tiering style and
diversity. In the absence of intervening event beds
interpretation of complex tiering can be difficult. In
Fig. 6f, a high diversity suite of mud/silt-filled traces
is associated with a silty to sandy substrate, represent-
ing moderate to low energy in a lower shoreface/
transition zone setting. Small-scale cross-lamination
may be evident, but this does not affect the tier
pattern. This is the most common type of tiering in
offshore transition facies in a genetically related
succession (e.g. Goldring et al., 1991).
Complex tiering in fine-grained sediments has been
extensively illustrated from chalks (Bromley and
Ekdale, 1986) and limestone micrite – marl cycles. It
has also been commonly seen in deep-sea sediments
such as hemipelagic deposits (Wetzel and Uchman,
1998a, 2001).
A difficulty that has been appreciated for some
years, and emphasised by Bromley (1996), is that as
aggradation proceeds the activity of deeper bioturba-
tors tends to obscure that of shallower bioturbators.
There are several situations where shallow tiers might
be advantageously preserved in multi-tiered sub-
strates:
(1) Below a distal event bed. This is the classic
situation for the preservation and casting of shallow
graphoglyptids on the soles of distal turbidites (Uch-
man, 1995) (Fig. 4).
(2) Below a storm event bed, where chance led to
minimal erosion.
(3) At an event bed, where the upper surface was
colonized by an infauna but, for various ecological
reasons, did not become established (Fig. 10c).
(4) In inclined heterolithic (alternating lithologies)
sediments, where minimal erosion took place with
each accreted layer. Gibert (1996) described this
situation from the Middle Jurassic marginal lime-
stones of central England. The description of Pleisto-
cene clinoform bedding in the Archakhangelos facies
of the Rhodes Formation on Rhodes, Greece (Brom-
ley and Asgaard, 1975; Hanken et al., 1996), and the
Middle Pleistocene Casarana Calcarenite of southern
Italy (D’Alessandro and Massari, 1997) probably
represent similar situations, but at a larger scale.
It is seldom that the traces of pioneers can be
observed intergrading into the equilibrium stage. A
diverse trace assemblage that can be shown to have
bioturbated the sediment contemporaneously will
almost certainly represent a climax association and
stable depositional environment, for example, tran-
sition zone and offshore facies comprising intensely
bioturbated muddy silts. It is generally agreed that
complex traces such as Zoophycos must have taken
a considerable time to be constructed, but the box-
works of Ophiomorpha, seen today in intertidal
settings, represent high activity in a dynamic envi-
ronment.
Tier modification is due to sequential environmen-
tal shift (below), associated with essentially continu-
ous sedimentation, though not necessarily at a unit-
form rate. In Fig. 6g, tier replacement, the initial tier
makers (opportunistic colonizers), were replaced by
different taxa, using different strategies. Bromley
(1996, Fig. 10.2-a) refers to this style of tiering, citing
examples from modern environments where each
change was due to an ecological factor (disaster),
and thus it is perhaps better interpreted as successive
attempts to colonize by different pioneers.
Gradational change in tier structure may be due
to deteriorating (Figs. 3 and 6i) or ameliorating
(Figs. 3 and 6h) conditions, referring to oxygen-
ation. Under deteriorating conditions burrows be-
come smaller, some are lost, and the deeper tiers
move upwards. This is well illustrated by deterio-
rating oxygenation (Wetzel, 1981; Wignall, 1994).
Deterioration may be sudden (Fig. 6i), or appear to
be sudden, for unrecognizable ecological factors,
leaving a frozen tier, for instance, below a deep-
water tephra event. In shallow water an erosional
sole to a tephra event is the general case and there
is no frozen tier (Goldring, 1999b).
Change in tier structure in fine-grained sediment
may be attributed to fluctuations in one or more
factors: oxygenation, food supply or sedimentation
rate (Wetzel and Uchman, 1998a). Intermediates
between the four ‘end-members’ shown by Wetzel
and Uchman (op. cit.) (Fig. 4) are to be expected. A
common feature of offshore transition zone– lower
 A. Taylor et al. / Earth-Science Reviews 60 (2003) 227–259 241

shoreface facies in the Fulmar Formation (Upper 5. Attributes of ichnofabric

Jurassic, UKCS) are alternations of a few mm
thickness between siltier lithologies with Phycosi-
phon or Planolites/Helminthopsis, to muddier lith-
ologies with Chondrites (e.g. Gowland, 1996; facies
association F).
Wetzel and Uchman (op. cit.) drew attention to a
modification to the trend depicted in Fig. 6h, during
slow re-oxygenation, which favoured the formation
of stabilizing microbial mats. Abrupt re-oxygenation
is probably due to bottom current activity associ-
ated with (slight) erosion. Decrease in burrow size
with depth only relates to functionally and behav-
iourally similar burrows (Wetzel and Uchman,
1998b).
4.1. Conclusions
The tier structure and sequential changes in tier
structure provide information:
(1) On ecological maturity, reflecting stability of
the environment;
(2) Highlights omission and hiatuses in sedimen-
tation, erosion or non-deposition;
(3) Reflects ecological changes, deterioration or
amelioration at and below the substrate surface due to
oxygenation, food supply, sedimentation rate, state of
substrate and other parameters;
(4) Provides information for the quantification of
porosity and permeability of the sediment;
(5) Appreciation of the effects of diagenesis and
consolidation of the sediment (Fig. 6j,k).
5.1. Primary sedimentology (lithofacies)
Every attempt should be made to document the
primary sedimentary characteristics of the bioturbated
rock type: lithology, grain size, grain sorting, scale
and type of preserved sedimentary structures (lithof-
acies).
5.2. Scaling the degree of bioturbation
To simply state that a sediment is bioturbated is
like saying that a sediment is fossiliferous: it does not
convey much of a visual impression. Furthermore,
categories such as low and high have little meaning in
that they involve too much subjectivity. Two schemes
for the assessment of bioturbation are in current use.
That of Taylor (in Taylor and Goldring, 1993), the
Bioturbation Index (BI) (Table 3), aims to relate the
degree of bioturbation to the preservation of the
primary bedding features. The technique requires a
little care but works well in terms of recognition and
reproducibility. The area (or length of core) classified
must be noted. Taylor’s scale is based on the Reineck
Scale developed through long experience of modern
sediments (Reineck, 1963). The scheme of Droser and
Bottjer (1987, 1989) is rather simpler, but only con-
siders the degree of bioturbation as the Ichnofabric
Index (ii). There is no zero, ii1 (Ichnofabric Index
One) indicating no bioturbation, and the scaling is
different.

Table 3
Bioturbation Index (BI) based on the amount of reworking with respect to the original sedimentary fabric (Taylor and Goldring, 1993)
Grade %Bioturbated Classification
0 0 No bioturbation

1 1–5 Sparse bioturbation: few discrete traces and/or escape structures

2 6 – 30 Low bioturbation: bedding distinct, low trace density, escape structures often common

3 31 – 60 Moderate bioturbation: bedding boundaries sharp, traces discrete

4 61 – 90 High bioturbation: bedding boundaries indistinct, high trace density with overlap common

5 91 – 99 Intense bioturbation: bedding completely disturbed, limited reworking due to repeated overprinting

6 100 Complete bioturbation: sediment reworking due to repeated overprinting

242 A. Taylor et al. / Earth-Science Reviews 60 (2003) 227–259

Fig. 7. Completely bioturbated sandy siltstone event beds from the Middle Jurassic Not Formation, Norwegian Continental Shelf. (A) Core
photograph of a Phycosiphon ichnofabric (dominant ichnotaxon), with a Bioturbation Index (BI) of 6, and an ichnodiversity of 5. (B) Schematic
diagram of the areal extent of the ichnotaxa present. (C) Ichnofabric Constituent Diagram of the Phycosiphon ichnofabric: on the horizontal axis
the percentage area of the sedimentary structures and burrow types plotted at log scale. On the vertical axis, events (in order) from the initial
sedimentation (primary fabrics—none preserved in this case) to subsequent modification by bioturbation (secondary fabrics).

The Bioturbation Index (BI) indicates the extent to
which the primary sedimentary structures are still
observable. Bromley (1996) has pointed out that
burrows may be reburrowed. This is covered by
BI = 6—overprinting, and it can be included in the
Ichnofabric Constituent Diagram (Fig. 7; Section 8)
(Taylor and Goldring, 1993) which plots the nature of
the primary sedimentation and the order of emplace-
ment and extent of the successive bioturbators.
5.3. Burrow size and frequency
Burrow size and frequency are aspects that need to
be addressed, especially when dealing with narrow
cores. While few burrows are of core diameter,
Thalassinoides can give an anomalous fabric, as can
pervasive Teichichnus. Sequential changes in burrow
size and frequency for individual ichnotaxa may be
due to ontogenetic factors and/or environmental shift.
The latter are frequently seen in tier modification.
Stunting has been attributed both to hypersaline con-
ditions (Gibert and Ekdale, 1999), and to low salinity
and an indication of estuarine environment. It is still
common for trace fossils with similar morphologies
but of different size classes to be assigned to different
ichnospecies. Stenomorphism (dwarfed; smaller than
typical) has not always been recognised as such
(Goldring et al., 1997), but substrate influence should
be considered: a common cause of the unusual shapes
of omission surface burrows and borings. Thalassi-
noides paradoxica reflects the increase in firmness of
the substrate restricting the degree of burrow exten-
sion previously available to the producer of T. suevica.
Size can also be a usefully considered factor in
discriminating between Taenidium ( < 10 mm breadth),
a regularly meniscate, back-filled burrow associated
with a marine ichnoassemblage, and Beaconites, char-
acteristic of many non-marine facies, which has an
irregular backfill reflecting passage of the animal
through heterolithic sediments (Goldring and Pollard,
1995). Large Beaconites (up to 25 cm breadth) prob-
ably represent vertebrate activity.
5.4. Ichnotaxonomy and ichnodiversity
Identification of the components of ichnofabrics
has two aspects: recognition of the mottling—the
biodeformational structures—and identifying the spe-
cific ichnotaxa. While the former has not yet in any
way been fully investigated, treatment of taxa in
vertical profile has been essentially geometrical (e.g.
in Ekdale et al., 1984). But when the Bioturbation
 A. Taylor et al. / Earth-Science Reviews 60 (2003) 227–259 243

Index is high, analysis is aggravated by reburrowing
and interpenetration, by cross-cutting, and compac-
tional effects (though burrows and borings made
contemporaneously tend to avoid each other). It is
not possible to identify, without question, a number of
ichnotaxa from a single section. And that is all that
may be available in a slabbed and resin embedded
core. This applies particularly to ichnotaxa such as
Diplocraterion and Teichichnus, or in the discrimina-
tion of Cylindrichnus, Asterosoma and Rosselia. The
likelihood of correct identification is increased with
repeated sequential occurrences in different cross-
sections. It would be advantageous if authors were
to indicate the degree of certainty for their identifica-
tion, at least in academic publications, by using
normal taxonomic procedure.
The question may be posed, whether it is essential
to take the trouble to name the components. The
question will certainly be debated, but an alternative
approach is possible along the lines of identifying
ichnoguilds (Section 6.2). Aspects that can confuse
identification include the quality of preservation
(taphoseries MacNaughton and Pickerill, 1995) (Fig.
8), and compactional aspects. These are of less con-
cern to taxa that can be identified in sections normal to
stratification.
The diversity of ichnotaxa associated with an
ichnofabric is an aspect often referred to in the
literature, especially that concerning brackish water
facies (e.g. Pemberton and Wightman, 1992), but is an
attribute that must be used with caution, and which is
rarely in itself of environmental significance. What is
of importance, however, is change in diversity
through a succession. Overprinting by key bioturba-
tors or deeper tier elite burrows may obscure shallow
tier burrows and reduce apparent diversity. However,
diversity may be overestimated (Jensen, 1997) when,
for instance, shafts (Skolithos) are counted independ-
ently of radiating traces (Chondrites) with which they
are linked functionally. Differences in preservation
have been cited frequently as separate taxa (Goldring
et al., 1997), and Uchman (1995) indeed favours the
use of ichnosubspecies to designate taphonomic dif-
ferences. Wetzel and Uchman (1998b) have shown
that biodeformational structures (Schäfer, 1956,
1972), which cannot be given ichnotaxonomic status
because of their obscurity, reflect bioturbation close to
the surface in soft sediment (essentially within the
mixed layer), and bioturbation at a deeper level. It is
in the latter that cross-cutting and overprinting (and
mechanical disruption) lead to destruction of earlier
burrows, but create new and identifiable traces as well

Fig. 8. A core-based example of a taphoseries (A) Ophiomorpha nodosa in characteristic mode with pellet-lined burrows. (B) Ophiomorpha
irregulaire, as irregularly compressed carbonaceous, mud-lined burrows. (C) Palaeophycus isp., as dense ramifications of weakly lined (often
top-lined) burrows. (D) Thalassinoides isp., as unlined burrows picked out by early chert cementation. Material from the Upper Jurassic Fulmar
Formation, UK Continental Shelf. Scale bars 5 cm.
 244
Table 4
Ichnofabric icon table for the Ophiomorpha ichnofabrics in Fig. 13

A. Taylor et al. / Earth-Science Reviews 60 (2003) 227–259

For each ichnofabric, a summary icon is drawn, and a definition given which incorporates sedimentological and ichnological parameters. Basic trace fossil data such as ichnotaxa,
diversity and degree of bioturbation (BI) is recorded along with colonization style. This type of table complements a lithofacies scheme and saves on extensive text generation.
 A. Taylor et al. / Earth-Science Reviews 60 (2003) 227–259
as biodeformational structures. Pollard et al. (1993)
appreciated the significance of mottling (i.e. biode-
formation) in discriminating between various Ophio-
morpha-dominated ichnofabrics (Table 4, Fig. 13),
and this should be extended further.
6. Aspects of ichnofabrics relating to a synthesis of
data
6.1. Trace fossils and ichnocoenoses, ecological
succession
In the 1970s, much effort was spent on attempts to
describe and determine palaeocommunities (reviewed
by Brenchley and Harper, 1998), in most cases from
shell beds and storm layers. In some cases trace fossils
were added to the lists of body fossils and included in
any palaeocommunity reconstruction (e.g. McKerrow,
1978). Where body fossils are absent, the trace fossil
assemblages have been interpreted as ichnocoenoses
in their own right. This has been successfully carried
out in respect of vertebrate footprints (Lockley et al.,
1994), who by neglecting lithology, incorrectly
referred the ichnocoenoses to ichnofacies. Three
aspects can be considered: (a) the trace fossil analysis
must be linked to the body fossil analysis, either as
part of a common autochthonous assemblage or,
indirectly, to a time-averaged assemblage; (b) the
tiering structure and ordering must be analysed to
distinguish between trace fossil assemblages of differ-
ent colonizations, e.g. pre- and post-event suites, (c)
care needs to be taken in measuring the diversity.
The ecological succession (ordering of the traces)
preserved in an ichnofabric can be determined from
the cross-cutting relationships and the tiering pattern
formed, for example, at the recolonization of event
beds, or at unconformities where repeated episodes of
erosion are frequently evident. Trace fossils that may
be interpreted as due to pioneers that speedily colo-
nize a vacant habitat, and opportunists generally,
display high abundance and small size (or burrow
diameter). This virtually implies high reproduction
and growth rates. A broad environmental tolerance
or adaptation to a stressed environment may be less
readily established, though occurrence in a range of
lithologies may be indicative of the former. But,
intensely bioturbated beds of a few centimetres thick-
245
ness due to pervasive deposit feeders (a single taxon)
are typical of non-marine floodplain and delta top
environments, e.g. Planolites montanus (Pollard,
1988).
6.2. Ichnoguilds
The guild concept (groups of species that exploit a
particular environmental resource) has proved useful
in ecology and has been extended to ichnology (e.g.
Bromley, 1996). As yet few studies have been carried
out. Potentially, ichnoguilds will form a useful basis
for universal palaeoenvironmental interpretation and
in the understanding of ichnofabrics. Each guild may
be defined on three categories: (a) the spatial relation-
ship i.e. tier level; (b) dwelling or semi-permanent
burrows OR vagile, transitory burrows; and (c) food
source: deposit feeding OR suspension feeding OR
farming (or chemosymbiosis). For example, in the
Upper Cretaceous Chalk of northwest Europe, and in
deeper basinal settings, the Chondrites– Zoophycos
ichnoguild can be recognised as deep-tier, non-vagile
deposit feeding (or chemosymbiotic) structures.
Besides Chondrites and Zoophycos, certain types of
Teichichnus, and also Trichichnus may be included.
These traces represent deep burrowing specialists.
7. Compaction and diagenetic factors
Compaction of trace fossils is, for the most part,
well understood, though aspects such as longitudinal
microfaulting associated with the compaction of mud-
filled burrows can be mistaken for bioglyphs. Diage-
netic effects are likewise well understood (Figs. 1 and
6). In ancient sediments it is seldom that tiering can be
readily interpreted in full. This is due not only to
cross-cutting and minor erosion but also to diagenetic
(and later metamorphic) processes. These lead to
enhancement (Fig. 6j), obliteration (Fig. 6k), or frag-
mentation of the ichnofabric. The relatively greater
permeability of the fill of once open burrows is latent
for preferential lithification (Fig. 6j) (elite trace fossils
of Bromley, 1996), though in other situations it may
be the matrix that is cemented (Fürsich, 1973b). In
extreme cases, this may lead to an open ‘plumbing
system’ such as may be seen in the Upper Cretaceous,
Chalk Rock (Bromley, 1967). More pervasive cemen-
246 A. Taylor et al. / Earth-Science Reviews 60 (2003) 227–259
tation through the whole sediment, or locally in
concretions, leads to reduced clarity of the ichnofabric
(Fig. 6k), but to advantageously preserved ichnofa-
brics in concretions in otherwise strongly compacted
mudrocks. Also, it is useful to mention the loss of
bed-junction traces with under- and over-beds. Com-
paction may also obscure the bioturbation (Fig. 1).
8. Presentation of ichnofabric data
Ichnofabric data can be readily incorporated onto
sedimentary logging sheets (Fig. 9). The trace fossils
can be symbolised and placed in the grain size profile
along with the sedimentary structures. The Bioturba-
tion Index (Table 3) can be plotted as a histogram
against depth and ichnodiversity recorded by interval.
The bracketing of the individual ichnofabrics will
reflect all of these parameters. Hiatal colonizations
(Fig. 10) may be coloured separately and stressed in
Fig. 9. Logging proforma for recording ichnofabric and bioclastic data.
the accompanying text. There is no formal agreement
as to the naming of ichnofabrics though it is gen-
erally convenient to use a group designation such as
‘Ophiomorpha/Palaeophycus ichnofabric’. Since the
study of ichnofabrics is still at an early stage it is
not surprising that, unlike the description of ichnof-
acies, different systems are being used in their desig-
nation, though all refer to the dominant ichnotaxa. But
it is illogical to select taxa that are not actually
present, as is frequently done with ichnofacies (Brom-
ley, 1996).
There are many pleasing graphical ways whereby
this information can be recorded and displayed. Some
of the alternatives include software packages such as
RAGWARE, STRATABUGS (biostratigraphical),
APPLECORE (sedimentological), and WELLCAD
(petrophysical), which allow the relative abundance
of individual ichnotaxa to be recorded by depth.
Before presenting summary diagrams, divisions
can be marked onto depositional models, and tier
 A. Taylor et al. / Earth-Science Reviews 60 (2003) 227–259 247

Fig. 10. Eight types of Key Stratal Surface and associated ichnofabrics. Modified and extended from Taylor and Gawthorpe (1993). Omission
with or without (or minimal) erosion. For details see text.

diagrams (Fig. 1) can illustrate the uniqueness of a 9. Ichnofabrics in stratigraphy

particular ichnofabric. The Ichnofabric Constituent
Diagram (Pollard et al., 1993) (Fig. 7) (Section 5.2)
provides a visual means for describing and comparing
ichnofabrics by: (1) assessing the extent and type of
the remaining primary fabric; (2) unravelling the
tiering and ordering of the ichnotaxa; and (3) recog-
nising the type of bioturbational event. The Ichnofabric
Constituent Diagram can be applied to hardgrounds
and rockgrounds (Goldring, 1995) that may reflect
changes in substrate consistency or water depth.
As trace fossils provide an in situ record of the
environment, and environmental changes, they lend
themselves to stratigraphic analysis. In effect, they are
excellent in defining the presence and nature of key
stratal surfaces, as well as contributing to the under-
standing of encasing facies. Ichnology can be success-
fully applied in both clastic and carbonate realms
because the trace fossils provide reliable data on the
nature of in situ sedimentary processes such as hiatus,
248 A. Taylor et al. / Earth-Science Reviews 60 (2003) 227–259
erosion, condensation, accumulation and exposure.
Recognition of the ‘surface’ is based on these prag-
matic observations rather than application of ‘univer-
sal ichnofacies’.
9.1. Ichnofabrics at key stratal surfaces
Ichnofabrics typically change at key stratal sur-
faces (Fig. 10) where omission is generally evident
and erosion common. Such surfaces are particularly
important in correlation, and in delimiting sequen-
ces and parasequences. The sedimentological pro-
cesses that may be involved are changes in flow
regime, omission, erosion (at all scales), consolida-
tion and lithification, subsidence, uplift, emersion
( F palaeosol formation). Each of these may be
reflected in the biotic recolonization by animals
and/or plants which typically accentuate the key
stratal surface.
Recolonization involves three distinct colonization
and tier-forming periods. Firstly, the pre-omission
suite (one of the several types of tier), which may
have been attenuated by erosion. Secondly, the omis-
sion suite, which colonized the substrate during omis-
sion or before sedimentation, was renewed. This suite
represents colonization that is generally associated
with increasing compaction, firmness and possible
lithification of the substrate, and may, rarely, show
an ecological succession (Gruszczynski, 1986).
Thirdly, the post-omission suite represents coloniza-
tion under renewed sedimentation. It may mark a
return to the pre-omission suite or to another style
of tiering. Subsequent burrowers will be influenced to
a greater or lesser extent by the (concealed) change in
substrate state at the omission surfaces. Changes
between the pre- and post-omission suites will gen-
erally be indicative of a significant shift in energy, and
frequently one that can be correlated with a change in
depth. Furthermore, the omission surface is frequently
complex due to more than one erosive event and there
may be a repetition of the omission- and post-omis-
sion suites, associated with greater attenuation of the
pre-omission suite.
Nine types of key stratal surface (Fig. 10) that are
associated with bioturbation are recognised. These fall
into two groups.
(1) Where no overall shift in environment has taken
place (Fig. 10a– e).
(a) A flooding event is indicated by Diplocraterion
within a non-marine (or lagoonal) succession. Knowl-
edge of marine, brackish, and non-marine ichnotaxa
and shelly fossils is necessary to recognise the sig-
nificance of the event, which is generally marked by
single tier (opportunist) colonization. The event may
be of short duration.
(b) Alternatively, rooted horizons on a delta flood-
plain may be cross-cut by a more diverse omission
suite of marine traces, suggesting the higher salinities
of an interdistributary bay, to be truncated, in turn, by
fluvial channel facies.
(c) In muddy, offshore sediments mid- to deep-tier
traces of the pre-omission suite (Teichichnus) are
cross-cut by shallow-tier burrows (Phycosiphon and
Planolites), and deep-tier Chondrites, generating a
hiatus overprint.
(d) Omission with change in substrate state but
without significant erosion or change in depth. Mid-
to deep-tier softground traces (Teichichnus, Chon-
drites) are shown crosscut by firmground traces (Sko-
olithos), and followed by further softground traces. A
likely situation is an offshore depositional environ-
ment: the omission surface representing a maximum
flooding surface or sediment by-pass. The degree of
lithification (firmground or hardground) will depend
on the lithology and duration of omission.
(e) In carbonate facies omission may be repre-
sented by a hardground. There is no significant differ-
ence in facies below or above the hardground. Rarely,
the ecological succession of trace fossils may indicate
gradual (submarine) lithification. Most commonly in
shallow water settings emersion was involved. The
correlative value of hardgrounds, especially where
geochemical factors were involved, is high (Gruszc-
zynski, 1998).
(2) Where a major shift in depositional environ-
ment has taken place (Fig. 10f – i).
(f) Omission with erosion leading to a change in
substrate state and a significant change in water depth
(energy state), but without emersion. Such exhumed
firmground surfaces, marked by vertical or sharp-
margined, passively filled burrows are typical of many
sequence boundaries, or combined sequence boun-
dary/transgressive surfaces. In the literature, the omis-
sion colonization is often referred to as Glossifungites
ichnofacies (e.g. in Pemberton, 1992). Such a surface
can also form at a major unconformity when lithifi-
 A. Taylor et al. / Earth-Science Reviews 60 (2003) 227–259
cation has not taken place, as at the Upper Cretaceous/
Tertiary unconformity in southern England (Bromley
and Goldring, 1992).
(g) Omission and emersion where a substantial fall
in relative sea level has taken place, and which is
generally accompanied by substantial erosion. A
rooted surface may develop under such conditions.
(h) Where relative deepening has taken place in a
marine environment (from shoreface to offshore),
there may be little or no evidence for any erosion
having taken place. But there will be a sharp change in
tier structure and, generally, of substrate type.
(i) Where a major change in facies has taken place
marking a significant flooding event, as by lagoonal
sediments transgressing floodplain sediments with
Beaconites. Again, little erosion may have taken
place. A precursor event may be evident just below
the key surface, which some would regard as marking
the formation boundary.
9.2. Ichnofabrics in genetically related successions
Between Key Stratal Surfaces the facies succes-
sions are essentially conformable and either show
little or no change, or display evidence for environ-
mental shift, as in upward coarsening (shallowing)
parasequences, representing a change from offshore
muds to shoreface sand (Fig. 11). Analysis of genet-
ically related successions provides the details of facies
interpretation and basin development. Other com-
monly observed changes in ichnofabric seen in genet-
ically related successions may be attributed to:
proximal –distal trends of turbidites and storm beds,
oxygenation trends in black shales and turbiditic
facies, salinity fluctuations in marginal marine (lagoo-
nal) facies, depth-related lateral changes in firm- and
hardgrounds.
Changes in the ichnofabrics may be gradual or
abrupt but display changes in primary sedimentary
structures, and other factors discussed above, that can
be correlated with the zonation of the shoreface and
offshore depositional environments. However, the
validity of inferences made on such changes within
genetically related successions has to be established
and this can only be achieved by observations made
on repeated occurrences in the field and in wells.
One problematic area is associated with recognis-
ing regression. For example, continued progradation
249
of a barrier shoreline will result in a rooted sandy
shoreface overlain by lagoonal mudstones. This is part
of a conformable genetic succession. However, enthu-
siastic stratigraphers may be tempted to place a
sequence boundary at the rooted surface (exposure
surface) and thus separate the genetically related
deposits. Ichnofabric analysis can provide useful
information on the palaeosol and aspects of both the
shoreline and lagoonal facies which may establish the
true nature of this environmental shift.
In practice, it is unlikely that any two successions
will display identical sedimentology and ichnology. If
the differences are considered to be significant, then
any depositional model has to take these into account.
For instance, in a depositional setting that is tectoni-
cally active, differences may be attributable to fault-
related mechanisms or salt diaparism leading to sedi-
ment by-passing or starvation. Such changes can be
recognised by departure from predicted genetically
related successions (Gowland, 1996).
As trace fossils record a biotic response to ambient
conditions, it is no surprise that individual depositio-
nal environments are often characterised by distinctive
styles of bioturbation. The application of ichnofabric
analysis in clastic deposits goes beyond merely cor-
roborating other sedimentological data. Trace fossils
commonly provide the sole line of evidence concern-
ing, for example, sedimentation rates or subtle envi-
ronmental shifts. Indeed, in highly bioturbated
deposits, primary sedimentological structures are
commonly obliterated. In such cases, palaeoenviron-
mental interpretation entirely rests on the successful
reading of the trace fossil succession, e.g. in Fulmar
Formation, United Kingdom Continental Shelf
(UKCS) (Martin and Pollard, 1996).
9.3. Ichnofabrics at event beds
Colonization and tiering may be interrupted by
event beds: storm beds, tempestites, turbidites, slump
deposits and tephras are the principal events. Event
beds are frequently composite and the record of
colonization of the older event(s) may have been
eroded. Perhaps the biggest difficulty facing geolo-
gists regarding muddy sediments, especially deep-sea
deposits, is in resolving the ichnology of the thin
( < 10 mm) mixed layer, especially the graphoglyptids
(Fig. 4). It is only on the soles of more distal
250 A. Taylor et al. / Earth-Science Reviews 60 (2003) 227–259

Fig. 11. Example of genetically related successions from the Upper Jurassic Franklin Sandstone Formation, UK Continental Shelf. Profiles from
a proximal and distal setting of a shoreline-shelf succession demonstrate systematic changes in ichnology and lithology in response to
progradation. Lateral changes in colonization style associated with key stratal surfaces are highlighted.

turbidites that these are preserved as pre-event
hypichnia, providing a ‘snapshot’ of the muddy sedi-
ment, and demonstrating the gentle erosive action of
the current.
9.3.1. Response to major ‘event’ sedimentation
Kern (1980) (see also Goldring, 1999a) recognised
criteria for distinguishing between the pre- and post-
turbidite suites of trace fossils, respectively, those
picked out by the turbidity current from the back-
ground mud and then infilled, and those which have
resulted from colonization from the top of the bed. In
most siliciclastic turbidites, it is rare to see burrows
traversing a bed because of the nature of the sediment.
If no erosion took place the pre-turbidite suite may
have been ‘frozen’, which is particularly important for
investigating the tiering in the mudrock. The tops of
turbidites are often intensely bioturbated, especially in
late Mesozoic and Caenozoic sediments, following
the rapid increase in diversity of the post-event suite at
the end of the Cretaceous, especially by echinoids
(Seilacher, 1977, 1986).
Although allowance must be made for unrealised
biological factors, six styles of event bed colonization
are recognised (Fig. 12).
(a) No colonization has taken place above an unfos-
siliferous or poorly fossiliferous (body fossil) event
bed. Absence of substrate recolonization in proximal
turbidites may be indicative of derivation from a non-
colonized fluvial/deltaic sand source, but may also give
an indication of the nature of the finer-grained suspen-
sion load. ‘Muddy’, graded beds, generally with
rippled tops, are typically not recolonized, probably
because of the unsuitability of the substrate for occu-
pancy. Such sediments appear to be most common in a
pro-delta setting due to a high sedimentation rate from
suspension flows (Walker and Plint, 1992).
(b) Animals (and plants) are often able to escape
from the smothering effects of an event bed by rela-
tively rapid escape. The degree of success varies within
 A. Taylor et al. / Earth-Science Reviews 60 (2003) 227–259 251

Fig. 12. Six styles of colonization at an event bed where an erosional base is normal. For details see text.

a population and between different groups. For modern
sediments, the evidence has been documented (Schä-
fer, 1972). This must be distinguished from tier adjust-
ment (Fig. 6c). The lower part shows bivalve
adjustment (based on Broadhurst et al., 1980) and the
upper part shows escape shafts (based on Goldring,
1964).
(c) Presence of a relocated biota, colonizing the top
of the bed, but without return to the pre-event biota.
Such a situation, for which we have no recorded
example, would suggest an appreciable change in
palaeogeographic conditions.
(d) Where colonization has led to a resumption of
the pre-event biota and bioturbation, with minimal
effects due to substrate change. This situation is
frequently encountered in association with storm
sheets of the order of 10– 100s m lateral extent (e.g.
Goldring et al., 1998).
We model two situations where recolonization was
relatively delayed (Fig. 12e,f). Sedimentation of the
upper part of the event bed, indicated here by a thick
(dm-m) tephra suggests initial recolonization by pio-
neers and opportunists. In Fig. 12e, initial coloniza-
tion was followed by ichnofabrics similar to the pre-
event suite. In the shoreface depositional zone this
more extensive disruption to benthonic life probably
relates to a situation following a tempestite, or tsu-
nami or widespread tephra event (Goldring, 1999b). If
the tephra event is widespread, the sole may well
constitute a key stratal surface. Large areas of the
seafloor were denuded of life by erosion and/or
blanketing by sediment.
252 A. Taylor et al. / Earth-Science Reviews 60 (2003) 227–259

In the second situation (Fig. 12f), the ichnofabric
above the event bed is shown as significantly different
from that in the pre-event sediments, but then returns
to this above. The ‘slow’ recolonization may lead to a
‘frozen’ tier ichnofabric followed by re-formation of
the pre-event ichnofabric, a situation typical of deep-
sea turbidites with relocation and/or recolonization
associated with the radical change in nature of the
substrate. But no change in overall palaeogeography
is indicated.
We are only now beginning to understand the
complexities of the processes of recolonization (Wet-
zel and Uchman, 2001). With thinner event beds,
particularly in more highly bioturbated Mesozoic –
Caenozoic sediments, many of the burrowers appear
to have survived the event by upward migration and
were able to relocate with the substrate surface. It is
only the shallower burrowers that recolonized.
Clearly, these scenarios are gradational to others.
10. Discussion
Each of the controls of ichnofabric character have
some environmental significance, but it is clearly the
manner of colonization and the tiering pattern, and the
changes in tiering that have taken place at key stratal
surfaces, at event beds, and through genetically related
successions, that are the most readily identified, and of
most significance for correlation and facies interpreta-
tion. The subtle changes in colonization and tiering that
may be observed in genetically related successions
provide high resolution in facies interpretation.
Trace fossils are currently being used in facies
analysis with two markedly different methodologies.
Both take into consideration the primary sedimentary
structures and other primary aspects of the sediment,
the processes involved and their interpretation. One
school (Pemberton, 1992) uses the specific ichnoge-
nera as indicators of ichnofacies, linked to an environ-
mental/bathymetric distribution model, the most recent
being that prepared by Frey et al. (1990). Frey et al.
(op. cit.) strongly argued that water depth is rarely the
governing factor for the distribution of trace fossils
and ichnofacies, and their diagram (op cit. Fig. 1)
shows several environmental locations of the Cruzi-
ana and other ichnofacies. The second school, for
example Martin and Pollard (1996) and Taylor and
Gawthorpe (1993), use the ichnofabric approach as
outlined here.
The advantages of using ichnofabrics in facies
interpretation have been detailed by Martin and Pol-
lard (1996) who showed that while three (Seilacher-
ian) ichnofacies can be recorded in the Upper Jurassic
Fulmar Formation of the North Sea, some 25 distinct
ichnofabrics can be recognised that are reflected in the
fine-scale environmental reconstruction achieved (see
also Fig. 13 and Table 4). This is because the
ichnofabric approach (as distinct from Seilacherian
ichnofacies) integrates all aspects of the sediment, and
thus reflects all of the parameters that control facies
formation and the aspect of key stratal surfaces.
However, Martin and Pollard (op. cit.) were con-
cerned almost wholly with the shore to offshore
profile. It is not proposed here to reiterate Bromley’s
(1996) critique of the ichnofacies approach, which
served sedimentology well for several decades. Fig.
11 gives an indication of the greater resolution that
can be achieved by using the ichnofabric method, and
Fig. 13 and Table 4 show that several ichnofabrics
associated with Ophiomorpha can be recognised, each
reflecting distinct environmental conditions.

Fig. 13. A range of Ophiomorpha ichnofabrics from Jurassic, Cretaceous and Eocene marginal marine, shoreface and deep water facies. (A)
Tidal channel. Medium to coarse-grained sandstones with Ophiomorpha and Planolites. Lower Cretaceous Woburn Sands, Chamberlain Barn
Quarry, Leighton Buzzard, England. (B) Tidal channel. Cross-bedded, well-sorted, fine-grained sands with woody debris, and O. nodosa shafts.
Upper Jurassic, Fulmar Formation, United Kingdom Continental Shelf (UKCS). (C) Washover sand. Slightly silty, fine-grained sandstones with
O. nodosa exhibiting floor deposits, and spreiten burrows. Upper Jurassic Fulmar Formation, UKCS. (D) Upper shoreface. Cross-bedded, fine-
grained sandstone with vertical shafts of O. nodosa. Middle Jurassic, Hugin Formation, Norwegian Offshore Continental Shelf (NOCS). (E)
Middle shoreface. Mottled, very fine-grained sandstone with Ophiomorpha, Palaeophycus and mottling. Upper Jurassic Fulmar Formation,
UKCS. (F) Lower shoreface. Very fine-grained, silty sandstones with O. nodosa and other traces. Eocene, Barton Formation, Barton-on-Sea,
southern England. (G) Lower shoreface with event beds. Alternation of sandy siltstones and silty, very fine-grained sandstones with O. nodosa,
Skolithos and Planolites. Miocene, Nigeria. (H) Sandy turbidite. Fine- to medium-grained, poorly sorted sandstone with Ophiomorpha. Upper
Cretaceous, Nise Formation, NOCS. (I) Debris flow. Medium-grained, poorly sorted sandstones with intraclasts and Ophiomorpha. Late
Palaeocene, Vaila Formation (UKCS).
 A. Taylor et al. / Earth-Science Reviews 60 (2003) 227–259 253

Seilacher (1967, 1978) referred to facies-breaking ichnotaxa exhibit different tiering styles in different
ichnotaxa, such as Chondrites and Planolites, that settings that can be used in the interpretation and
were not specific to any of his ichnofacies. But such distinction of facies as, for example, in the size and
254 A. Taylor et al. / Earth-Science Reviews 60 (2003) 227–259
penetration depth of Chondrites (Fig. 3). The environ-
mental tolerances that may be applied to several
ichnotaxa are still in dispute.
A literature search by Bottjer et al. (1987, 1988)
indicated that there were environmental shifts through
time in the distribution of Zoophycos, and its gradual
restriction to slope and basinal settings in the later
Mesozoic, and to Ophiomorpha, which extended its
distribution into deep-water sediments in the Meso-
zoic. Skolithos, or ‘pipe-rock’, becomes relatively
uncommon in post-Palaeozoic sediments (Droser,
1991, 1993). Ichnological changes at the Precam-
brian – Cambrian boundary (Orr, 2001) may reflect
changes in the nature of the substrate, especially in
the role played by biomats, a factor not discussed by
Orr (op. cit.). An aspect of ichnofabrics that has not
yet been researched is stratigraphic differences due to
changes in tier structure and guild composition. These
may be due to evolutionary changes and/or to latitu-
dinal and geographic (climatic belt) changes and
emphasised by taphonomic processes. Most ichnotaxa
are long-ranging, and hence by implication, there are
relatively few ichnofabrics, when compared with body
fossils. Such changes in ichnofabric would be analo-
gous to changes in Mesozoic benthic shelf (shelly)
communities (for example, see Boucot, 1975; Aber-
han, 1994). Pemberton et al. (1992, Table 1) referred
to the distinctions between early Cretaceous, late
Cretaceous and Tertiary suites of trace fossils in
non-marine environments, which may be readily
attributed to the evolution of angiosperms, insects
and crustaceans. Ecological factors such as climate
may be responsible for the distribution of Ophiomor-
pha, as presently understood, or the apparent absence
of typical Diplocraterion from post-Miocene sedi-
ments (except for the relatively diminutive burrows
of Corophium and Polydora), though known world-
wide from Cambrian to Miocene strata. Perhaps the
producer of Diplocraterion is known from modern
tropical deltas but not its burrow.
Significant geographic differences can be illus-
trated by the distribution of some modern trace-form-
ing taxa. For instance, living limulids are unknown
along the eastern Atlantic seaboard and interpretation
of European Mesozoic xiphosurid traces (e.g. Wang,
1993) is much influenced by North American docu-
mentation (e.g. Caster, 1938). While the J-form bur-
row of Arenicola sp. is characteristic of northwest
European beaches, the enteropneust Balanoglossus
sp. and the holothurian Leptosynapta sp., forming
U-burrows, are characteristic of the southeastern coast
of the USA. The bioturbational effects of Arenicola
and Balanoglossus are essentially similar and repre-
sent two parallel communities of Thorson (1957).
Latitudinal variation in bioturbation has recently
been considered by Cadée (2001) who, in particular,
emphasised the lack of deep-burrowing callianassid
crustaceans in arctic and temperate coasts. The broad
distinction between carbonate sediments found in
tropical as distinct from cool-water environments is
often referred to as that between the chlorozoan/
chloralgal and foramol facies, respectively (Lees,
1975). This can probably be extended to the ichnol-
ogy of Cretaceous and Cenozoic carbonate sediments.
Ichnofabrics of tropical stratiform carbonate sedi-
ments are dominated by deep-tier traces formed by
arthropods, whereas in cool-water shelfal carbonates,
shallower tiers, especially those due to the activity of
spatangoid echinoids, are advantageously preserved.
Problems arise especially with the elucidation of
the ichnology of marginal marine facies where the
ichnotaxa are frequently assigned to a mixed Skoli-
thos – Cruziana ichnofacies. Pemberton (1998) and
Gingras et al. (1999) indicated six ichnological criteria
for the recognition of brackish water facies (modify-
ing earlier work of Pemberton and Wightman, 1992):
a low diversity of trace forms and suites dominated by
a single ichnogenus, overall diminutive traces, simple
marine forms constructed by trophic generalists, ver-
tical and horizontal traces common to the Skolithos
and Cruziana ichnofacies, and locally prolific trace
densities. In our experience of estuarine/delta distribu-
tary facies, and applying the attributes of ichnofabrics
discussed here, we would modify these to emphasise
the opportunistic nature of colonization in this envi-
ronment; the often insufficient time that was available
to a constructor to form a burrow system, or even a
lamination-destroying mottling. Thus, burrows may
be poorly formed and ‘incomplete’ and the primary
lamination distinct (BI 1 –2). Diversity may be low
especially in relation to an open marine situation.
Under tropical, monsoonal conditions strong outflow
of fresh water and mud may extend the effects far out
onto the prodelta shelf necessitating care in facies
interpretation. It is probable that most emphasis here
must be placed on the primary sedimentary attributes.
 A. Taylor et al. / Earth-Science Reviews 60 (2003) 227–259
We do not recognise burrow size per se as a distinctive
attribute, since Ophiomorpha is so characteristic of
Mesozoic – Caenozoic situations and is generally of
‘normal’ size. In wind weathered profiles, slender (1
mm diameter) traces may weather out (e.g. Pollard et
al., 1993, Fig. 4D), but are invisible on slabbed core,
and are similar to those mentioned by Gingras et al.
(1999, p. 372) from an upper estuarine situation.
Perhaps the telling sentence regarding diminution is:
‘‘there is some question as to the nature and mecha-
nisms by which diminution (commonly observed in
these deposits) occurs. We suggest this is a reflection
of population composition as opposed to true dimin-
ution’’ (Gingras et al., 1999, p. 372).
The question may be posed as to whether the
recognition and analysis of colonization style and
tiering patterns in ichnofabrics might replace their
characterization by ichnotaxa (essentially by pattern
recognition), despite the difficulties in understanding
taphonomy, ichnological and zoological taxonomy,
and establishing satisfactory ichnotaxobases (Goldr-
ing et al., 1997). The answer must be in the negative,
since facies interpretation is best achieved by taking
as fully an integrated approach as possible, though
identification is best accompanied by taphonomic and
ethological interpretation (Goldring, 1999a; Goldring
et al., 1997) and also because of the environmentally
specific nature of many ichnotaxa, especially the
depth-specific nature of many borings.
However, many ichnotaxa are in practice (see
above) identified on only partial morphology—as
with resin-mounted and slabbed core. Indeed, the full
morphology of many is unknown or known only in
part. For instance, Cylindrichnus is often applied to a
burrow cross-section, and Ophiomorpha to a fragment
of shaft or gallery. This may be sufficient for those
relying on an identification alone but, since trace
fossil morphology is so much influenced by tapho-
nomic and sedimentological factors, much valuable
information may be lost by doing so. Nevertheless,
ichnoguilds (above) are likely to play an increasing
role in ichnofabric interpretation.
11. Conclusions
(1) Analysis of both the sedimentological and
ecological parameters of bioturbation at all scales
255
provides a basis for the description and interpretation
of an ichnofabric.
(2) Colonization and tiering are the most signifi-
cant factors in ichnofabric analysis. Eleven styles of
tiering and their controls are recognised.
(3) General models for tier organization in fine-
and coarser-grained sediments are developed. The
mixed layer is only well-formed in fine grained sedi-
ments, while the transition layer can be anything from
a few millimetres to several metres thick.
(4) There are several methods by which tiering can
be identified in ancient sediments, and four situations
favour the preservation of shallow-tier traces: below a
distal event bed, below and immediately above an
event bed, and on inclined heterolithic sediments.
(5) Many traces are difficult to identify in slabbed
cores or vertical profiles. This can lead to errors in
facies interpretation.
(6) Two categories of biodeformational structures
(mottling: where identification to ichnotaxon is not
possible) are important: those that are produced in the
mixed zone (or very shallow tier), and those formed at
deeper levels by cross-cutting. Discrimination
between types of the latter could prove useful in
ichnofabric analysis.
(7) Greater recognition and emphasis of ichno-
guilds might be more useful in evolutionary studies
of ichnofabrics than emphasising ichnotaxa.
(8) The Colonization Window, the period of time
which is available for successful colonization of the
substrate, is extended to include chemical (oxygen,
salinity), and trophic (bottom food availability) as
well as physical (wave, current energy) and biological
(spatfall) ‘windows’.
(9) Event beds (storm beds, turbidites, etc.) affect
the tiering. Recolonization may take place with only
minor interruption, and a return to the pre-existing
trace-producing biota, or the substrate may remain
uncolonized, or recolonization may take place by an
entirely new suite of organisms.
(10) The ichnology at Key Stratal Surfaces can be
used to understand relative sea level changes associ-
ated with basin development, hiatus, omission and
sediment bypass.
(11) Description and analysis of the ichnology of
Genetically Related Successions is placed on a firm
basis for the integrated interpretation of basin devel-
opment.
256 A. Taylor et al. / Earth-Science Reviews 60 (2003) 227–259
(12) Integrated ichnofabric analysis provides far
greater resolution than does ichnofacies analysis in the
interpretation of ancient sedimentary environments.
(13) The ongoing debate on the ichnology of
marginal marine, especially estuarine, facies is not
resolved, but greater emphasis must be placed on the
primary sedimentology and the Colonization Window.
(14) An integrated approach to ichnofabric analy-
sis provides for better insight into subsequent dia-
genesis of the sediment and porosity vs. permeability
relationships.
Acknowledgements
We are indebted to many petroleum companies and
their staff, including Amarada Hess, Norsk Hydro,
Phillips, Saudi Aramco, Shell, Statoil, TotalFinaElf,
for the opportunities we have been given, professio-
nally and academically, to examine cores and well
logs. We wish to thank John Pollard (University of
Manchester), as postgraduate supervisor and friend to
AMT, and to him and Gerhard Cadée (Netherlands
Institute for Sea Research, NIOZ) for their advice and
criticism of this contribution. We also wish to thank
Andrew Thurlow and Chris Elliott (Ichron) for their
help in detailed ichnological data collection.
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Andrew Taylor commenced work on a trace
fossil-based PhD in 1988 at the University
of Manchester, England, under the guid-
ance of John Pollard. Here, he worked on
the ichnology, facies analysis and sequence
stratigraphy of North Sea Middle Jurassic
fluvio-deltaic strata. This was followed by
4 years of post-doctoral research at Man-
chester (sponsored by Norsk Hydro) on the
stratigraphic analysis of bioturbated shal-
low-marine reservoir analogues in the Ter-
tiary of the Spanish Pyrenees. In 1995, he joined the geoscience
consultancy Ichron, where he is principally involved in play fairway
analysis, field-scale stratigraphic modelling and running training
courses in ichnology and stratigraphy.
Roland Goldring, now retired from a Read-
ership at the Postgraduate Research Institute
for Sedimentology, The University of Read-
ing, has been interested in trace fossils since
his PhD days on the palaeontology and
stratigraphy of Upper Devonian marine
sediments of North Devon, England. This
was under the supervision of Scott Simp-
son, a student of Rudolf Richter. He appre-
ciated that the trace fossils provided much
insight into the interpretation of the deposi-
tional environment of ancient facies, and during the past decade,
following retirement, has become increasingly involved in the full
integration of sedimentology and ichnology: ichnofabrics. In collab-
oration with Ichron he has contributed to many ‘hands-on’ short
courses, on ichnology and ichnofabrics.
Stuart Gowland first became interested in
trace fossils whilst undertaking a PhD at
the University of Hull, England, under the
supervision of Martin Brasier. From this
point onwards he quickly realised the
potential of using trace fossils to refine
sedimentological interpretations and pro-
vide useful information for stratigraphic
correlation. He worked as a reservoir geol-
ogist for Geochem Group for 12 years,
prior to joining the international geosci-
ence consultancy group Ichron in 1995. Currently, he works on the
reservoir quality assessment of bioturbated oil and gas reservoirs
worldwide.