Ichnology Research Group The original work and research in this report was undertaken and/or collated

by Dr. A.J.Pulham & Prof. S.G.Pemberton. Unless otherwise indicated

copyright is retained by the authours. December 1998.

ICHNOLOGY ATLAS
Version 1.2 (May 1999)
Inspired by, and dedicated to the memory of, Robert W. Frey (1938-1992)
We stand on the shoulders of giants so we might better understand and see with greater clarity

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Ichnogenera Index The original work and research in this report was undertaken and/or collated
by Dr. A.J.Pulham & Prof. S.G.Pemberton. Unless otherwise indicated
copyright is retained by the authours. December 1998.

Anconichnus Helminthopsis Scolicia

Arenicolites Lockeia Skolithos

Asterosoma Macaronichnus Subphyllochorda

Bergaueria Ophiomorpha Teichichnus

Chondrites Palaeophycus Terebellina

Conichnus Planolites Thalassinoides

Cylindrichnus Rhizocorallium Zoophycos

Diplocraterion Rosselia
Gyrolithes Schaubcylindrichnus
 ANCONICHNUS
Description: Irregularly meandering, black-cored burrows with a
pale halo of coarser silt. In cross-section, the burrows are elliptical
to sub-circular, U-shaped in the longitudinal profile. Burrow
structure is normally erratic, vermicularly tangled (Bromley,
1990), and often short in length. Extensive bioturbation tends to
create a mottled fabric, obscuring most traces (Goldring et al.,
1991). This mottled fabric is often overlooked due to overprinting
by deeper tier organisms. In core, Anconichnus commonly
appears as tiny dark pin-head sized spots (transverse section) or
dark lines (longitudinal section), which may be discontinuous and
surrounded by a pale silt halo.

Interpretation: Anconichnus represents the grazing trails of

worm-like organisms. Goldring et al., (1991) suggests the halo
surrounding the fecal core is due to parapoidal sorting of the
sediment by the trace-making organism. Pemberton (pers. comm,
1991) indicates the halo may be the effect of oxidation by oxygen-
rich water introduced into the substrate by the organism’s burrow.
In this case, Anconichnus may be equivalent to Helminthopsis.

Trophic Classification: Systematic feeding/grazing.

Environmental Considerations: Anconichnus is a common

element of the Cruziana and Skolithos ichnofacies on a normal
marine shallow shelf and is often found as an opportunistic
colonizer of storm-deposited sands. Anconichnus is associated
with Chondrites, Terebellina, Planolites and other shallow tier
trace fossils.

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ARENICOLITES
Description: Simple, vertical, U-shaped tube with no spreiten between the limbs. Exterior walls
generally smooth with no ornamentation; apertures of one or both tubes may flare. Generally preserved
in full relief, but may be recognized in plain view by paired openings.

Interpretation: Arenicolites is interpreted as the dwelling burrow of an annelid or a small crustacean.

Probably originators include the polychaete Arenicola or the amphipod Corophium. A predominate
suspension feeding tropic mode has been postulated for the organism dwelling in such structures.

Trophic Classification: Suspension- or filter-feeding.

Environmental Considerations: Arenicolites is generally associated with arenaceous substrates in low

energy shoreface or sandy tidal flats. A common element of the Skolithos ichnofacies. When found in
great numbers it can be indicative of mixed tidal flats.
Modern
Tracemakers
Arenicola: modern burrow Arenicola: modern lugworm

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 ASTEROSOMA
Description: Star-shaped burrow system consisting of radial bulbous arms tapering inwards towards an
elevated centre. The arms tend to be circular in cross-section and consist of concentric lamination of sand
and clay packed around a central tube; the exterior is generally smooth, but may exhibit longitudinal striae
or wrinkles.

Interpretation: Based on the tubular construction of galleries and the details of sediment working,
Asterosoma has been interpreted as the feeding burrow of a worm. The organism seems to have probed
repeatedly into the sediment to enlarge the gallery and work more sediment vertically and laterally; exact
details of this process remain conjectural. The sediment fill may be related to feeding/waste stowage
functions.

Trophic Classification: Selective deposit-feeding by a vermiform organism.

Environmental Considerations: Asterosoma represents a specialized feeding structure and is therefore

more commonly associated with fully marine conditions. Generally found in the upper lower shoreface
associated with Rosselia. A common form of the Cruziana ichnofacies.

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 BERGAUERIA
Description: Cylindrical to hemispherical, plug-shaped, vertical burrows
with smooth, unornamented walls; circular to elliptical in cross-section;
infillings essentially structureless; rounded base with or without shallow
central depression and ridial ridges. The length-diameter ratio generally
varies from 2:2 to 2:8.

Interpretation: Bergaueria represents the activities of actinian anemones.

Ethologically, two interpretations have been postulated, in which it
represents a resting trace (cubichnia) or dwelling burrow (domichnia).
Both are probably correct, with lined specimens (e.g. B. langi) representing
domichnia and unlined specimens (in most cases) representing cubichnia.

Trophic Classification: Most actinian sea anemones can be classified as

carnivores.

Environmental Considerations: Generally Bergaueria is indicative of

normal marine conditions on a wave- or tide-dominated shoreface. It is a
common element in the Skolithos ichnofacies. Found in brackish-water
environments but greatly reduced in size.

Modern Tracemaker: Sea Anemone

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 CHONDRITES
Description: Chondrites is a complex root-like burrow system of regularly branching feeding tunnels of uniform
diameter which never anastamose, interpenetrate, nor cut across one another (photo taxis). Branching typically is
in the form of side branches (up to five or six orders) angling off a higher order or main tunnel at 30 degrees to 40
degrees rather than bifurcating at Y-shaped junctions.
In core, Chondrites commonly appears as an array of tiny elliptical dots where the vertical slice through the core
truncates the numerous branching tunnels. In some instances, longitudinal sections through individual tunnels
and broken portions of branches are exposed.

Interpretation: It has been suggested that Chondrites represents tunnels produced by a deposit-feeding
sipunculid, which worked from a fixed centre on the substrate surface and created tunnels by extending its
proboscis. However, some forms of Chondrites penetrate so deeply into the substrate that they could have been
only produced by a vermiform animal dwelling within the structure, moving bodily through the sediment in the
manner of the modern polychaete Heteromastus.

Trophic Classification: Chondrites represents a complex deposit-feeding strategy.

Environmental Considerations: Although a well known facies crossing form Chondrites is common element of
the Cruziana ichnofacies. It represents a complex feeding behaviour and is therefore more commonly associated
with fully marine conditions. A monospecific association of Chondrites has been interpreted to be indicative of
low oxygen zones. Possible tracemaker: Sipunculid

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CONICHNUS
Description: Conical, amphora-like, or acuminated
subcylindrical structures oriented perpendicular to
bedding; base may be rounded or may exhibit a
distinct papilla-like protuberance. Fillings may reveal
patterned internal structures such as chevron laminae
but not radial medusoid symmetry. The lining, albeit
very thin, constitutes a distinct discontinuity between
the infill and the adjacent matrix and is often subject to
diagenetic altercation.

Interpretation: Conichnus has been interpreted as

representing domichnia, cubichnia, or anemone-like
organisms (Frey and Howard, 1981). Two
ichnospecies are currently recognized, C. conicus and
C. papillatus. Criteria for their recognition are based
primarily on slight differences in the overall burrow
outline and the presence or absence of a distinct apical
protuberance.

Trophic Classification: Anemones tend to be

carnivorous.

Environmental Considerations: Conichnus is

generally associated with higher energy, arenaceous
middle shoreface environments deposited under
normal marine conditions. Commonly associated with
the Skolithos ichnofacies.
Possible tracemaker:
Sea Anemone

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 CYLINDRICHNUS
Description: Cylindrical, subcylindrical to subconical
burrows, straight to gently curved, having multiple
concentrically layered walls. Orientations range from vertical
to horizontal; never branched.

Interpretation: The most diagnostic feature of this form is

its multilining which has been attributed to the activity of the
organism in its burrow, responding to a slow, continuous
sedimentation rate; as sediment entered the burrow, it was
pressed against the burrow wall. Multi-lined burrow walls
have been recognized in structures produced by the
polychaete of the genus Nereis, or the crustacean genus
Callinassa. Although predominantly vertical systems have
been attributed to suspension-feeding crustaceans, horizontal
systems have been attributed to deposit-feeding worm-like
organisms.

Trophic Classification: Suspension-feeding dwelling

burrow.

Environmental Considerations: Cylindrichnus is a common

element of the Skolithos ichnofacies and the proximal end of
the Cruziana ichnofacies. Also commonly associated with
sandy tidal flats and a common element (with Skolithos) in
lateral accretion deposits within estuarine channels.

Modern Nereid burrow

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DIPLOCRATERION
Description: Vertical, U-shaped spreiten burrows. Spreiten may be
retrusive, protrusive, or a combination of both (see Vol. I, p. 15, Fig. 9);
apertures of the tubes may be cylindrical or funnel-shaped; limbs of U may
be parallel or divergent. In some instances may appear in core as dumbbell-
shaped burrows on the top of units; the paired circular openings are joined
by a horizontal band of reworked sediment corresponding to the spreite.

Interpretation: The systematics of Diplocraterion and other vertical, U-

shaped ichnogenera were reviewed by F rsich (1974a). Based on an
analysis of morphological features, F rsich interpreted Diplocraterion as
the dwelling burrow of a suspension-feeding organism. Probably
originators include polychaetes, echiuroids, and crustaceans (amphipods).

Trophic Classification: Suspension-feeder.

Environmental Considerations: Diplocraterion is a common element in

the distal end of the Skolithos ichnofacies in middle shoreface settings. It is
also common on sandy tidal flats and in estuarine channel deposits.

Possible tracemaker:
the Amphipod Corophium

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 Possible tracemaker:
the Polychaete Heteromastus

GYROLITHES
Description: Dextrally or sinistrally coiled burrows up to several centimeters high. Whorls are several
millimeters in diameter and are generally isodiametric. In core, Gyrolithes appears as layers of paired tunnels
converging upwards or downwards.

Interpretation: A similar spiral burrow is created by the capetellid polychaete (Notomastus).

Trophic Classification: Burrow represents the dwelling burrow of a suspension-feeding organism.

Environmental Considerations: Gyrolithes occurs in wave-dominated bays, splay sands, and has a tolerance of
very low salinity environments. Commonly found in a high density monospecific assemblage.

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HELMINTHOIDA / HELMINTHOPSIS
Description: Regular meandering (Helminthoida), or irregularly meandering (Helminthopsis), smooth-walled
burrows which never branch, interpenetrate or cut across one another. In cross-section, the burrows are elliptical to
sub-circular and are generally horizontal. Fill tends to be dissimilar from the surrounding matrix. In core,
Helminthoida/Helminthopsis commonly appears as tiny dark spots (transverse section) 1 - 3 mm in diameter, or dark
lines (longitudinal sections).

Interpretation: Helminthoida/Helminthopsis represents the grazing trails of worm-like organisms.

Trophic Classification: Systematic grazers.

Environmental Consideration: Helminthopsis is a common element of the distal Cruziana ichnofacies a

proximal Zoophycos ichnofacies on a normal marine shallow shelf. In some instances, Helminthopsis can be
associated with low energy, fine-grained bay environments.

Possible tracemaker: Nematode

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 LOCKEIA
Description: Small almond-shaped oblong bodies preserved in convex hypo
relief. Mostly symmetrical with smooth unornamented surfaces tapering to
sharp and obtuse points at both ends. Dimensions are highly variable with
lengths varying from 2 to 12 mm.

Interpretation: Lockeia represents the resting trace (cubichnia) of a bivalve.

The pit is created by extending the foot.

Trophic Classification: Lockeia could represent the activities of both

suspension feeding and deposit feeding bivalves.

Environmental Consideration: Lockeia is best preserved in environments

subjected to frequent episodic depositional events. It is common in tidally-
influenced zones such as tidal point bars and tidal flats. Found in brackish-
water and fresh-water environments.

Possible tracemaker:
bivalve foot
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MACARONICHNUS
Description: Unlined, distinctly walled, predominantly horizontal,
randomly curving and meandering to distinctly spiraled cylindrical
burrows that rarely interpenetrate, never branch, and commonly
possess a thin mantle concentration of mafic mineral grains; infills
markedly cleaner than host sand and are either structureless or
indistinctly backfilled.

Interpretation: Based on the modular morphology and the nature

and composition of the burrow fill, Macaronichnus is interpreted as
the feeding burrow of an opheliid polychaete (Clifton and
Thompson, 1978).

Trophic Classification: Selective deposit-feeding by a vermiform

organism.

Environmental Considerations: Macaronichnus represents a very

specialized adaptation to a stringent set of environmental parameters.
This trace fossil occupies high energy foreshore to proximal upper
shoreface environments (Saunders and Pemberton, 1986). It is also
present in some high energy tidal shoal units associated with zones at
the mouth of a large estuary (Wach and Ruffell, 1991).

Possible tracemmaker: the polychaete Euzonus

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 Possible tracemaker:
Thalassinid shrimp

OPHIOMORPHA
Description: Simple to complex burrow systems distinctly lined with agglutinated pelletoidal sediment. Burrow
lining is near-smooth on the interior; densely to strongly mammalated or nodose on the exterior. Individual
pellets or pellet masses may be discoid, ovoid, conical , mastoid, bilobate, or irregular in shape. Characteristics of
the lining may vary within a similar composition to host rock but in some instances it may be filled actively with
meniscate laminae. Branching is irregular and, where present, Y-shaped; at bifurcations, burrows become
swollen.

Interpretation: The origin and significance of the morphological features exhibited by Ophiomorpha were
discussed by Frey et al., (1978). Based mainly on the character of the pelletal burrow lining, they recognized four
ichnospecies: O. borneensis, O. irregularire, O. nodosa, and O. annulata. In well-burrowed offshore sediments,
wall linings are thin and poorly developed and the species of the ichnogenus is somewhat intergradational with
Thalassinoides. Ophiomorpha represents the dwelling burrows of decapod crustaceans, including numerous
species of Thalassinidean shrimp.

Trophic Classification: Dwelling burrow of suspension-feeding shrimp.

Environmental Considerations: Commonly associated with the Skolithos ichnofacies, prolific numbers in
marine shoreface environments. Also found in brackish water, sandy substrates including estuaries and tidal
shoals.

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 Possible tracemaker: bamboo worm

PALAEOPHYCUS
Description: Infrequently branched, distinctly lined, cylindrical, horizontal to inclined burrows in which the
sediment fill typically is of the same lithology and texture as the host stratum. Wall linings may be smooth,
longitudinal striated or with transverse annuli.

Interpretation: Palaeophycus is distinguished from the morphologically similar ichnogenus Planolites

primarily by wall linings and the character of burrow fills (Pemberton and Frey, 1982). Fills of Palaeophycus
represent passive, gravity-induced sedimentation within open, lined burrows; the fillings therefore tend to be of
the same composition as the surrounding matrix. Passively filled, lined burrows are typically interpreted as
dwelling structures. The predaceous glycerid polychaete, Glycera, has been taken as an excellent modern analog
for the Palaeophycus organism.

Trophic Classification: Dwelling burrow of predaceous polychaete.

Environmental Considerations: Associated with the Skolithos ichnofacies in both high energy and low energy
shoreface environments. Commonly found with Planolites or Macaronichnus; can also be found in episodic
storm sands and brackish-water assemblages.

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PLANOLITES
Description: Unlined, rarely branches, straight to tortuous, smooth to
irregularly walled or annulated burrows, circular to elliptical in cross-
section, of variable dimensions and configurations; fillings essentially
structureless, differing in lithology from host rock.

Interpretation: Planolites is distinguished from Palaeophycus primarily

by having unlined walls and burrow fills differing in texture from that of the
adjacent rock; fills may differ in fabric, composition, and colour as well.
Fills of Planolites represent sediment processed by the tracemaker,
especially through deposit-feeding activities of mobile endobients (bottom-
feeders), probably infaunal polychaetes or other worm-like organisms.

Trophic Classification: Feeding burrow of deposit-feeder.

Environmental Considerations: Found in virtually all environments from

freshwater to deep marine.

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RHIZOCORALLIUM
Description: Straight to sinuous, horizontal, U-shaped
spreiten-burrows. Tubes are generally distinct and more or
less parallel; ratio of tube diameter to diameter of spreite is
1:5. Spreite are typically protrusive. Burrow infill is
commonly identical to the matrix, but in some cases is
finer-grained. In core, Rhizocorallium is discerned by two
circular burrows (tube arms) joined by a horizontal band
(spreite). Preserved primarily as endichnia.

Interpretation: F rsich (1974b) discussed the systematics,

morphology, and ethology of Rhizocorallium. Based on
major and accessory features, he divided Rhizocorallium
into three ichnospecies: R. jenese, representing more or
less straight, short, and commonly oblique spreiten burrows
interpreted as the domichnia of suspension feeders;
R. irregulaire, representing long, sinuous, branching and
planispiral forms; and R. uliarense, representing
trochospiral spreiten burrows. The latter two are interpreted
as fodinichnia of deposit feeders. Although originally
interpreted as a coral, a sponge, or an alga, the presence of
distinct scratch marks on the tube wall is consistent with a
crustacean origin.

Trophic Classification: Dwelling/feeding burrow of a

deposit-feeder.

Environmental Considerations: Generally associated

with fully marine environments from the distal Cruziana
ichnofacies. Under certain conditions it may occur in the
Glossifungites ichnofacies.

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 ROSSELIA
Description: Single entrance, cylindrical, vertical to inclined, straight or
gently curved burrows with an opening expanded to a funnel shape. The
bulb is filled with finer grained sediment in poorly developed concentric
layers. In transverse section, the bulbs are circular to sub circular and attain
greatest diameter at the bedding surface. The cylindrical, central shaft
displays concentric layering, is commonly gently curved, and in some cases
penetrates the bulb.

Interpretation: The basal stem of Rosselia typically is identical in

construction with that of Cylindrichnus. Chamberlain (1971) and others
have noted fabricational similarity between Asterosoma and Rosselia, and
Frey and Howard (1982) reported occasional but complete intergradations
among specimens individually referable to Skolithos, Cylindrichnus,
Rosselia, and Asterosoma. Two ichnospecies of Rosselia are currently
known: R. socialis and R. rotatus, separation of ichnospecies is based on
the nature of the infill of the expanded opening. Rosselia is interpreted as a
feeding burrow of polychaete.

Trophic Classification: Feeding/dwelling burrow of a deposit feeder.

Environmental Consideration: Generally associated with the proximal

Cruziana ichnofacies in full marine settings. When found along or with
Asterosoma, it is a good indicator of the top of the lower shoreface.

Possible tracemaker: Nereid polychaete

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 Possible tracemaker: Sabellearid worm

SCHAUBCYLINDRICHNUS
Description: Plural curving tubes, gently arcuate with upper ends of the tubes being vertical, whereas lower
ends are approximately horizontal. Isolated sheaves or bundles of closely juxtaposed, congruent, well lined,
equisize tubes that do not taper, branch, or interconnect. Number of tubes making up an individual sheaf is
variable but as many as 20 per sheaf have been described. Both interior and exterior surfaces of the tubes are
smooth and unornamented.

Interpretation: Schaubcylindrichnus is interpreted as the communal domicile of either suspension feeders or

head-down tubicolous deposit feeders such as the modern maldanid polychaete Clymenella torquata.

Trophic Classification: Suspension-feeders or tubicolous deposit feeders.

Environmental Considerations: Schaubcylindrichnus is most common in shoreface settings, slightly seaward

of true high-energy parts of the strand but well landward of quiet, low-energy parts of the offshore zone.
Schaubcylindrichnus has not been reported from brackish water deposit.

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SKOLITHOS
Description: Single entrance, cylindrical to subcylindrical, straight to
curved, vertical to sub vertical, unbranched burrows that do not cross over
or interpenetrate. The shafts are either lined or unlined with generally
smooth walls, but may be annulated. The infill is typically structureless.

Interpretation: Ethologically, Skolithos represents the dwelling burrows of

suspension-feeding organisms or passive carnivore. A multitude of
probable originators have been postulated, including: the polychaetes
Sabelleria, Arenicola and Onuphis, the phoronid Phoronopsis and insect
larvae.

Trophic Classification: Dwelling burrow of suspension-feeding vermiform

organism.

Environmental Considerations: Lined specimens generally associated

with marine or brackish environments. It is an element of the Skolithos
ichnofacies. But because Skolithos can be constructed by many different
kinds of organisms it is found in virtually every type of environment from
marine to non-marine.

Possible tracemaker: Burrow of modern

the polychaete Onuphus polychaete Longonereis
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SUBPHYLLOCHORDA
(SCOLICIA)
Description: Subphyllochorda has the appearance in core of a series of vertical to sub horizontal, concave-up
laminae. On a plan view, not seen in core, there can be well-developed medial ridges, with striae slanting back
from the middle. Subphyllochorda has the appearance in split core of a large Teichichnus. It can be differentiated
by the relatively large size and more singular occurrence compared to the smaller and more densely distributed
Teichichnus.

Interpretation: Subphyllochorda is formed by the forward migration combined with passive and active backfill
by the feeding organism. Scolicia is produced by echinoids and gastropods. Echinocardium cordatum are
recorded in Pleistocene sediments (Bromley and Asgaard, 1975), at the ends of their backfilled burrows, with their
spines still in active digging position.

Trophic Classification: Feeding (fodinichnia) burrow of a deposit-feeding organism.

Environmental Consideration: Wide-ranging, found from the lower shoreface through offshore to deep-water
slope environments within the Cruziana, Zoophycos and Nereites ichnofacies and is also recorded from flysch
deposits.

Possible tracemaker:
the heart urchin
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 Possible tracemaker: polychaete worms

TEICHICHNUS
Description: Teichichnus appears in split-core sections as a vertical series of
tightly packed concave-up or (more rarely) concave-down, crescentric laminae.
Longitudinal sections show wavy, long laminae that usually merge upwards at the
ends. It is formed by the upward migration of a horizontal to sub horizontal tunnel
produced by an organism moving back and forth in the same vertical plane, probing
the sediment for food.

Interpretation: The Teichichnus - producing animal appears to be a deposit-

feeding, worm-like organism which migrated upward in its burrow to keep pace
with sedimentation.

Trophic Classification: Dwelling/feeding burrow of a deposit-feeding organism.

Environmental Consideration: Commonly found in lower shoreface to offshore

environments associated with the Cruziana ichnofacies. Prevalent in lagoon/bay
facies characterized by brackish-water. Never associated with freshwater.

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TEREBELLINA
Description: Subcylindrical, vertical, gently to strongly curved burrows
with circular to elliptical cross-sections. Diameters are highly variable
and the tubes gradually taper distally. The lining is very distinct, ranges
in thickness fro 1.5 to 5 mm, is composed of either calcium carbonate or
sand grains, and is more resistant to weathering than either the
surrounding matrix or the burrow fill. Material infilling the burrow is
similar in composition to the host rock. In core, specimensgenerally
appear to be horizontal due to the curvature of the tube.

Interpretation: Terebellina was originally interpreted as the body fossil

of a polychaete; however, because it is a tube constructed by the
organism, it must be considered an ichnofossl. Based on its
morphological characteristics, Terebellina can be interpreted as the
dwelling burrow of a suspension-feeding polychaete. A single valid
ichnospecies, T. palachei, is known.

Trophic Classification: Dwelling burrow of suspension-

feedingorganism.

Environmental Consideration: Commonly found in the distal Cruziana

ichnofacies in offshore, marine environments. Special adaptation for
constructing an open tube in argillaceous substrates. Also found in fine-
grained, brackish-water bay envionments.

Possible tracemaker: Sabeallarid polychaetes

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 Possible tracemaker: Upogebid shrimp

THALASSINOIDES
Description: Relatively large burrow systems consisting of smooth-walled cylindrical components. Branches are
Y- to T-shaped and are enlarged at points of bifurcation. Burrow dimensions may vary within a given system and
cross sections range from cylindrical, half-moon shaped to elliptical. Most systems are essentially horizontal with
some irregularly inclined.

Interpretation: Very thinly-lined to essentially unlined burrow systems are characteristic of fine-grained
coherent substrates, in which wall reinforcement is unnecessary. Structureless to parallel-laminated or graded
burrow fills represent passive (gravity-induced) sedimentation, whereas meniscate or chevron-laminated
sediments represent active backfilling by the tracemaker. Thalassinoides is generally regarded as a dwelling
and/or feeding burrow of a decapod crustacean (Thalassinid shrimp). Enlarged junction points are often used as
turning points for the organism, or as breeding chambers.

Trophic Classification: Dwelling/feeding burrow of a deposit-feeding crustacean.

Environmental Consideration: Associated with the Cruziana ichnofacies in lower shoreface to offshore
environments. Also found in low diversity, brackish-water suites.

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ZOOPHYCOS
Description: Zoophycos is basically a circular to lobate sheet-like spreite,
either flat, curved, inclined or wound in screw fashion around a central
vertical axis. The spreite is a horizontal or sub horizontal web of closely
juxtaposed parallel burrow tunnels. Each tunnel in the burrow system
presumably represents the path of its feeding apparatus during a single
probing of the sediment. Successive probing side-by-side in the same plane
produce a horizontal spreite.

Interpretation: Ekdale (1977) interpreted Zoophycos to be the feeding (or

grazing) structure produced by a vermiform organism with a fully extensible-
retractable body (as in the phylum Sipunculids). Other interpretations
suggest that the tracemaker was an annelid.

Trophic Classification: Grazing trace of a deposit-feeding organism.

Environmental Consideration: Associated with the distal Cruziana and

Zoophycos ichnofacies in fully marine, offshore shelf environments.

Possible tracemaker: Echurid

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RC
University of Colorado at Boulder

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