INTEGR. COMP. BIOL.

, 43:219–228 (2003)

The Proterozoic and Earliest Cambrian Trace Fossil Record; Patterns, Problems
and Perspectives1

SÖREN JENSEN2
Department of Earth Sciences, University of California, Riverside, California 92521

SYNOPSIS. The increase in trace fossil diversity across the Neoproterozoic-Cambrian boundary often is
presented in terms of tabulations of ichnogenera. However, a clearer picture of the increase in diversity and
complexity can be reached by combining trace fossils into broad groups defined both on morphology and
interpretation. This also focuses attention on looking for similarites between Neoproterozoic and Cambrian
trace fossils. Siliciclastic sediments of the Neoproterozoic preserve elongate tubular organisms and structures
of probable algal origin, many of which are very similar to trace fossils. Such enigmatic structures include
Palaeopascichnus and Yelovichnus, previously thought to be trace fossils in the form of tight meanders.

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A preliminary two or tripartite terminal Neoproterozoic trace fossil zonation can be be recognized. Pos-
sibly the earliest trace fossils are short unbranched forms, probably younger than about 560 Ma. Typical
Neoproterozoic trace fossils are unbranched and essentially horizontal forms found associated with diverse
assemblages of Ediacaran organisms. In sections younger than about 550 Ma a modest increase in trace
fossil diversity occurs, including the appearance of rare three-dimensional burrow systems (treptichnids),
and traces with a three-lobed lower surfaces.

INTRODUCTION shaped burrows (Bergaueria, Conichnus) (Narbonne et

There was an increase in the diversity and com-
plexity of trace fossils across the Neproterozoic-Cam-
brian boundary (Seilacher, 1956). Broadly speaking,
Neoproterozoic trace fossils are typically horizontal,
unbranched trails or burrows made close to the sedi-
ment surface. In the Cambrian, animals probed deeper
into the sediment and produced more complex bur-
rows, including branching forms, with a concomitant
expansion in the range of sizes. The bulk of the Neo-
proterozoic–earliest Cambrian trace fossil record
comes from siliciclastic strata deposited in shallow
subtidal settings. Also other environments are repre-
sented including evidence for colonization of the con-
tinental slope (e.g., Crimes, 2001). Examination of
sections world wide led to the realization that a rea-
sonably consistent order exists in the first order of ap-
pearance of several trace fossil types, which permitted
erection of trace fossil based zones (Crimes, 1987,
1992, 1994; Narbonne et al., 1987; Walter et al.,
1989). Based on successions in Newfoundland, Nar-
bonne et al. (1987) recognized three trace fossil zones
straddling the Neoproterozoic-Cambrian boundary—
the Harlaniella podolica, Treptichnus pedum, and Ru-
sophycus avalonensis zones—each matching a global
trace fossil zone erected by Crimes (1987). The Trep-
tichnus pedum Zone is characterized by the first ap-
pearance of an assemblage of trace fossils including
complex three-dimensional burrows (e.g., Treptichnus
pedum), the first arthropod-type scratch marks
(Monomorphichnus), and several ichnotaxa of plug-
1 From the Symposium The Cambrian Explosion: Putting the
Pieces Together presented at the Annual Meeting of the Society for
Integrative and Comparative Biology, 2–6 January 2002, at Ana-
heim, California.
2 Present address: Area de Paleontologia, Facultad de Ciencias,
Universidad de Extremadura, 06071 Badajoz, Spain; E-mail: soren@
unex.es
al., 1987; Narbonne and Myrow, 1988). It was even-
tually decided to select the base of the Treptichnus
pedum Zone at a section at Fortune Head, Burin Pen-
insula, Newfoundland, as the basal Cambrian GSSP
(Brasier et al., 1994; Landing, 1994). Subsequent stud-
ies have corroborated and expanded the broad-scale
stratigraphic application of the trace fossil based zo-
nation in the Lower Cambrian (e.g., MacNaughton and
Narbonne, 1999), although there has been some mod-
ification in the stratigraphic ranges of some key taxa
at the Burin Peninsula stratotype section (Gehling et
al., 2001).
Trace fossils not only are useful in earliest Cambrian
stratigraphy, but their importance has been recognized
in addressing questions related to the appearance of
animals and the interpretation of the Cambrian ‘explo-
sion’ (Seilacher, 1956; Bergström, 1990; Conway
Morris, 1993, 1998; Valentine, 1995; Erwin, 1999).
The importance of early trace fossils is particularly
great if one favors the view that key bilaterian features
could only have developed in a moderately large ben-
thic animal (e.g., Valentine, 1994, 1995; Budd and Jen-
sen, 2000; Collins and Valentine, 2001). There have
been numerous reports of trace fossils older than 600
Ma, but upon critical evaluation these traces have all
proven to be misidentified inorganic structures or me-
taphytes, or have been misdated (e.g., Crimes, 1994;
Hofmann, 1992; Fedonkin and Runnegar, 1992; Run-
negar, 1992a, b). Prominent recent reports of trace fos-
sils more than 1 Ga BP (Seilacher et al., 1998; Ras-
mussen et al., 2002) also are questionable (Conway
Morris, 2002; Budd and Jensen, 2003). In fact, all un-
doubted trace fossils postdate the Marinoan (or Var-
angerian) Ice Age, which ended about 590–570 Ma
BP.
The purpose of this paper is to briefly examine the
Neoproterozoic trace fossil record both in terms of its

219
220 SÖREN JENSEN

broad patterns and its relation to the Cambrian trace

fossil record.
CAVEATS OF TRACE FOSSIL NOMENCLATURE
Discussions of Neoproterozoic and Cambrian trace
fossils have largely been conducted at the level of
ichnogenera or ichnospecies. Indeed, compilations of
trace fossil diversity largely consist of tabulations of
ichnotaxa. Naming of invertebrate trace fossils is
largely based on morphology, and carries no implica-
tion of a commonality of trace producer (e.g., Mag-
wood, 1992; Pickerill, 1994; Bromley, 1996). A prob-
lematic aspect of trace fossil naming is that more often
than not, only a limited portion of a trace fossil is

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visible or preserved. Furthermore, the same trace pre-
served at different levels, such as preservation at the
interface of two contrasting types of sediment, can re-
sult in morphologically distinct traces. What is in ef-
fect a single biogenic structure therefore can be as-
signed to different ichnogenera depending on what
portion of the trace is visible (or preserved) to the
observer; this is particularly problematic for forms that
are three-dimensionally complex. It can be argued that
naming each trace purely by its preserved morphology
is the most objective approach, and that subjective
lumping may lead to loss of information. Nevertheless,
particularly for the study trace fossils at the Neopro-
terozoic–Cambrian, I find it preferable to look for sim-
ilarities, and to critically evaluate the taphonomy of
each morphological feature. A benefit of this approach
is that it better allows attempts to look for links be-
tween Neoproterozoic and Cambrian trace fossils.
Below, selected Neoproterozoic trace fossils will
therefore be discussed under broad groupings, defined
partly by criteria of morphology but which also incor-
porate significant interpretative elements (cf., Zhu,
1997; Jensen et al., 2000).
NEOPROTEROZOIC TRACE FOSSILS
Horizontal unbranched trace fossils
The archetypical Neoproterozoic trace fossils are
unbranched irregularly meandering to sinuous burrows
and trails typically a few millimeters in width, prob-
ably formed within less than 10 mm to the sediment-
water interface (Fig. 1). Depending on their general
pattern, these traces have been assigned to the ichno-
genera Planolites, Helminthoidichnites, Gordia, Coch-
lichnus, Helminthopsis and Helminthoida (e.g., Hof-
mann, 1990; Narbonne and Aitken, 1990). In Neopro-
terozoic examples of these ichnotaxa, traces can typi-
cally be followed along bedding planes or cleavage
surfaces for less than a few tens of centimeters, com-
plicating ichnogeneric assignment. The Neoprotero-
zoic Helminthoidichnites, Helminthopsis and Gordia
are all broadly comparable in that they represent tran-
sient movement through the sediment, perhaps as feed-
ing/scavenging traces. These traces are found in sandy
sediments and appear to have been formed as tempo-
rary tunnels as the animal forced itself through the
sediment. The subsequent fate of the burrow depended
FIG. 1. Simple trace fossils on the upper surface of a sandstone
bed. Neoproterozoic, South Australia. Specimen is in collection of
James G. Gehling, Adelaide. Scale bar is 10 mm.
on the properties of the sediment above and below the
tunnel, and microbial binding may have played an im-
portant role in preservation (Gehling, 1999). Speci-
mens preserved in negative relief on the sole of beds
demonstrate that this type of trace formed within the
sediment (Seilacher, 1999), but probably close to the
sediment water interface. Bedding planes may be
densely covered with these traces, often giving the im-
pression of true branching. In no case can it be proven
that these are not created by intersection of two traces
or by secondary successive branching (cf.,
D’Alessandro and Bromley, 1987), where an animal
has followed the course of an older trace. In principle,
all of these forms may have been created by the same
type of animal. The absence of ornamentation (i.e.,
scratch marks) must be understood as a preservational
artefact and does not imply original absence of such
structures.
Of particular interest is the occurrence of rare mod-
erately regular meanders (Narbonne and Aitken,
1990). These can be included in Helminthorhaphe
(sensu Uchman, 1995) though they do not have the
closely guided meander systems of some Phanerozoic
forms. In the Ediacara Member in South Australia,
Helminthorhaphe occurs sparsely on surfaces densely
covered by Helminthoidichnites and shares the same
style of preservation. These guided meanders are not
spurious because an exceptionally preserved specimen
from the Flinders Ranges contains a Helminthorhaphe,
which grades into an involute/evolute spiral (Runne-
 PROTEROZOIC TRACE FOSSILS
FIG. 2. A. Schematic drawing of a Helminthorhaphe continuous
with a Spirorhaphe from the Ediacara Member, Flinders Ranges,
South Australia, compared with the type material of Spirorhaphe
involuta. The Neoproterozoic specimen was examined and traced in
the field, and is figured by Runnegar (1992a, fig. 3.9). B. Morpho-
logical change in a trace fossil such as Archaeonassa as a function
of position of animal relative to sediment water interface. White is
sand, dark shading mud. Cross-section of animal (light shading) is
hypothetical. Compare with Figure 3B.
gar, 1992a). The spiral is similar in shape and size to
the holotype of Spirorhaphe involuta (Stefani, 1895)
from the lower Tertiary of Karpathos, Greece (Fig.
2A). The next oldest occurrence of Spirorhaphe is
from the Ordovician of New Brunswick (Pickerill,
1980).
Dwelling burrows
The case for Neoproterozoic permanent or semi-per-
manent dwelling structures is problematic. Neoproter-
ozoic globular to plug-shaped forms variously called
Beltanelliformis, Beltanelloides, Nemiana and Hage-
netta often show similarity with Phanerozoic reports
of Bergaueria (Fedonkin and Runnegar, 1992; Crimes,
1994). The latter is generally interpreted as a cnidarian
burrow, formed by sea anemones (e.g., Alpert, 1973).
The Neoproterozoic forms likely are body fossils rath-
er than trace fossils, though this distinction may be
difficult to recognize in practise (cf., Crimes and Fe-
donkin, 1996).
The case for Neoproterozoic dwelling burrows of
bilaterian origin is problematic. Glaessner (1984, p.
66), compared short grooves with pointed ends from
the Ediacara Member of South Australia to the base
of U-shaped dwelling traces. These, however, are pre-
served in negative hyporelief with no additional ex-
tension into the sediment and therefore more likely are
external molds of an organism (unpublished observa-
tion). Narbonne and Aitken (1990) figured specimens
of Palaeophycus tubularis from the Mackenzie Moun-
221
tains which are reasonable candidates for broad, flat
U-shaped burrows, which may have been at least semi-
permanently occupied. A similar observation can be
made for some small traces often referred to Planolites
ballandus and Planolites montanus (see Walter et al.,
1989).
There appears, however, to be no Neoproterozoic
evidence for accommodation to sediment influx, re-
sulting in vertical sediment stacking (spreite). Also, the
case for Neoproterozoic Skolithos (Crimes and Germs,
1982; Fedonkin, 1985) is doubtful. For example,
Crimes and Fedonkin (1996) now consider that the
Namibian specimens proposed to be Skolithos more
likely represent body fossils. Grazhdankin and Ivant-
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sov (1996, p. 677) maintained the trace fossil identi-
fication of Skolithos declinatus, inclined structures
from the late Vendian of the White Sea area (Fedonkin,
1985), reporting densities of 245 burrows/dm2. More
detailed information on this interesting form is needed.
Horizontal trace fossils with levees; Archaeonassa
and Psammichnites
An animal that pushes itself through the upper por-
tion of a sandy layer will displace sediment to either
side. The morphology of this type of trace fossil will
depend on how deeply the animal is submerged in the
sediment (Fig. 2B), the sediment properties, as well as
morphology of the animal. In sediments with high ten-
sile strength, displaced sediment may rupture to form
segmented levees (cf., Knox and Miller, 1985). Neo-
proterozoic trace fossils of this type have been report-
ed as Aulichnites and Nereites (Narbonne and Aitken,
1990; Crimes and Germs, 1982; Jenkins, 1995), but
they can all be included in the broadly defined Ar-
chaeonassa (cf.,Yochelson and Fedonkin, 1997). Re-
ports of Neoproterozoic Nereites (Crimes and Germs,
1982; Jenkins, 1995) appear to be based on material
in which the segmented lobes formed through rupture
of sediment with higher tensile strength (cf., Knox and
Miller, 1985). By contrast, the lateral lobes of Phan-
erozoic Nereites are constructional, formed by repeat-
ed probing in the sediment (e.g., Orr and Pickerill,
1996).
A slab from the Ust Pinega Formation, along the
White Sea (Fig. 3A) shows several specimens of Ar-
chaeonassa that have a wide flat central area flanked
by relatively narrow raised lobes. Other specimens
demonstrate the great importance of trace morphology
as a function of trace depth relative to the sand surface
(Fig. 3B).
The Neoproterozoic Archaeonassa are of particular
interest in that they provide probable links to the Low-
er Cambrian ‘‘Taphrhelminthopsis’’ circularis (Fig. 4).
Seilacher (1997), Zhu (1997) and Seilacher-Drexler
and Seilacher (1999) explicitly, or implicitly, included
the lower Paleozoic reports of Taphrhelminthopsis in
Psammichnites, interpreting it as a collapsed top sur-
face above the actual burrow. In the model of Seilacher
(1997) these are internally complex trace fossils that
are morphologically distinct, depending on what part
222
FIG. 3. A. Archaeonassa isp. from the Ust Pinega Formation, winter coast of the White Sea, north-west Russia. (Sedgwick Museum, Cam-
bridge, SM 27518). Positive epirelief. Scale bar is 10 mm. B. Several morphologic varieties of Archaeonassa isp. in the Ediacara Member,
Flinders Ranges, South Australia, reflecting depth of animal movement within the sediment (specimen in collection of J.G. Gehling, Adelaide).
Positive epirelief. Scale bar is 10 mm.
of the trace fossil is exposed. However, most Lower
Cambrian ‘‘Taphrhelminthopsis’’ circularis have been
described only based on upper surface morphology
with no information on internal structures (but see
Hofmann and Patel, 1989; Hagadorn et al., 2000). Re-
lated Lower Cambrian forms include Protospiralich-
nus, Planispiralichnus and Qipanshanichnus (Crimes
et al., 1977; Fedonkin, 1985; Luo et al., 1994).
Valentine (1995) hypothesized that most horizontal
Neoproterozoic trace fossils were produced by mol-
lusk-like animals. The flat surface between the levees
in Archaeonassa from northern Russia (Fig. 3A), cer-
tainly is consistent with such a producer. Conway Mor-
ris and Peel (1995) derived molluscs and annelids from
halkieriids or their ancestors. The body fossil Kimber-
ella has recently been re-interpreted as a mollusc-like
animal (Fedonkin and Waggoner, 1997). There has
even been reports of Kimberella found in association
with structures interpreted as radular scratch marks
(Seilacher, 1997; Fedonkin, 2001). There have been no
detailed documentation of the nature of this interesting
FIG. 4. Field photograph of ‘‘Taphrhelminthopsis’’ circularis on
upper surface of sandstone bed. Lower Cambrian, Chapel Island
Formation, Burin Peninsula, Newfoundland. Scale bar is 10 mm.
SÖREN JENSEN
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association and the purported radular marks, but hal-
kierids or proto-halkierids make one attractive search
pattern for the producer of Archaeonassa-type trace
fossils.
PROBLEMATIC ‘‘TRACE FOSSILS’’
Purported Neoproterozoic tightly guided meanders
and fecal rows
Somewhat surprisingly there are reports of tightly
guided Neoproterozoic meanders (e.g., Crimes and Fe-
donkin, 1994). Such trace fossils contrast to models
that depict evolutionary optimization of meander trac-
es during the Phanerozoic with increasingly effective
coverage starting from rather simple Cambrian speci-
mens (Seilacher, 1974). Subsequent findings (see
Crimes and Fedonkin, 1996) have shown that relative-
ly closely guided meanders were present also in the
Cambrian, but these are more loose than the Neopro-
terozoic forms. It now is clear that this interpretation
of Neoproterozoic forms should be abandoned and that
they probably are not trace fossils at all (e.g., Jensen,
1996; Seilacher, 1998; Gehling et al., 2000).
The most widely reported of these purported me-
ander traces is Palaeopascichnus Palij, 1976, consist-
ing of closely positioned wide or short, elongate bars
or grooves arranged in straight or winding rows (Fig.
5B). In most cases the width of the segments remain
about the same. Palaeopascichnus has a wide distri-
bution including Russia (Fedonkin, 1981), the Ukraine
(e.g., Palij, 1976), South Australia (Glaessner, 1969;
Jenkins, 1995), Poland (Paczesna, 1986), and New-
foundland (Narbonne et al., 1987; Gehling et al.,
2000).
Palij (1976) compared Palaeopascichnus to the larg-
er Phanerozoic meander Helminthoida but noticed that
Palaeopascichnus differs in that no turning points
could be observed. Indeed, Fedonkin (e.g., 1981) has
suggested that it was caused by an animal with a feed-
ing organ that operated transversely to the animals’
direction of movement.
Subsequently, ‘‘classic’’ meanders from the Neopro-
terozoic were reported as Yelovichnus gracilis (Fedon-
kin, 1985). On closer examination, however, there ap-
 PROTEROZOIC TRACE FOSSILS 223

FIG. 5. Palaeopascichnus and other problematic Neoproterozoic structures. Scale bar in all is 2 mm. A. Yelovichnus-type meander-like
preservation. Notice closed loop near base. Hypichnial view. Winter Coast of the White Sea, northern Russia Ust-Pinegia Formation (bed 1)
(SM 327517). B. Palaeopascichnus-type preservation. Notice ring at left of center where pelletoid is missing. Hypichnial view. Locality as

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in A, (SM 327516). C, D. The ‘metaphyte’ Orbisiana simplex from the Neoproterozoic of the Kunevichi-4 core, depth 557–558 m, western
Russia. Same specimen in normal photography and x-ray radiograph (D). The specimens occur in a greenish gray silty clay. Up/down orientation
not known. (Tallinn Institute of Geology, IG Va1841).

pear to be no examples in which the meander limbs
are actually connected by turning points. A typical
specimen (Fig. 5A) consists of closely juxtaposed flat-
tened oval structures with narrow raised rims. Al-
though not always apparent, in well preserved areas
these rims form closed ovals (Fig. 5B). The apparent
meanders are evidently an artifact arising from close
juxtaposition of the oval units. A similar effect of
raised rims occurs in Palaeopascichnus in the places
where the elongate bars are missing (Fig. 5B). Iden-
tical rimmed units also are found scattered on sediment
surfaces. Notably the same range of structures occur
in Neoproterozoic strata of both the White Sea area
and in the Ediacara Member of South Australia (cf.,
Jenkins, 1995, pl. 1d,i, pl. 2c). It therefore seems that
Palaeopascichnus and Yelovichnus are related, and re-
flect different preservational styles of the same struc-
ture.
A further likely preservational variety of Palaeo-
pascichnus occurs as low-relief elongate depressions
in calcisiltites of the Wonoka Formation in South Aus-
tralia (Haines, 2000).
Neoproterozoic simple and complex rows of pellet-
shaped objects that have been assigned to Neonereites
and interpreted as fecal pellets (e.g., Fedonkin, 1981),
should also be compared to Palaeopascichnus. Impor-
tant to this argument is the ‘metaphyte’ Orbisiana sim-
plex Sokolov, 1976, from the Neoproterozoic of Rus-
sia. Specimens from the Neoproterozoic of the Kunev-
ichi-4 core near Ladoga, western Russia, consist of
rows or aggregates of objects delineated by a pyritic
rim (Fig. 5C, D). There is a range of organization of
the form, from biserial strings forming long chains to
aggregates with an irregular outline. The exact shape
of the objects are not clear at the present, but they
appear to consist of aggregated spherical or hemispher-
ical bodies. Size differs between specimens but ap-
pears to remain more or less fixed within a specimen
(Fig. 5C, D). In the examined samples there are larger
specimens with a diameter of spheres 0.4–0.9 mm, and
smaller specimens with a diameter of 0.2–0.3 mm.
Sediment within the spheres appears to be identical to
that surrounding them. Forms that appear identical to
Orbisiana have also been described by Chistyakov et
al. (1984) from Neoproterozoic outcrops near lake
Onega, western Russia. They noted that strands may
be sequentially subdivided up to five times and consist
of elements about 1 mm wide (Chistyakov et al., 1984;
fig. 3B).
There is a strong resemblance between Orbisiana
simplex and the various Neoproterozoic forms reported
as Neonereites. For example, in Neonereites biserialis,
figured by Fedonkin (1981, Pl. 14, 1–5; 1994, Fig. 6A)
the sediment filled objects appear to have a narrow
sediment-free zone separating each other. Orbisiana
simplex with pyritized walls occurs in green fine silt-
stone, whereas Neonereites-type forms are found in
fine sandstone, and so the two may be different pres-
ervational aspects of the same structure.
An assignment of these objects is problematic.
Haines (2000) suggested that Palaeopascichnus from
the Wonoka Formation should be compared with
brown algae such as Padina rather than to trace fossils,
and Runnegar (1995) suggested that they are stratiform
stromatolites. Probably related, at least in terms of ta-
phonomy, are Mesoproterozoic bedding plane mark-
ings from the Belt Supergroup, USA and the Bange-
mall Group, Australia, which have been interpreted as
probable megascopic algae (Grey and Williams, 1990;
Horodyski, 1993; for further discussion of these forms
see also Fedonkin and Yochelson, 2002). Grey and
Williams (1990) reconstructed these as hollow or fluid-
filled spheres with toughened walls of probable algal
affinity. A similar interpretation seem likely for Pa-
laeopascichnus and Yelovichnus. More recently Leguta
and Seilacher (2001) suggested that these are xeno-
phyophoran protists.
For the purpose of this paper, exact affinities are not
at issue, but it is clear that a trace fossil origin must
be abandoned for Palaeopascichnus. Fedonkin’s
(1981) suggestion of vertical probes may need further
consideration for certain forms, but the apparent gra-
dation among the objects discussed above strongly ar-
224
FIG. 6. A. Lower surface of finely laminated siltstone/claystone with Harlaniella (top) and tubular fossils, superficially similar to trace fossils.
Scale bar is 5 mm. Pasha core, western Russia, Kotlin Stage (IG Va1843). Insets show areas magnified in B and C. B. Detail of lower right
portion of tubular fossil in A, with signs of brittle fracture. Scale bar is 2 mm. C. Detail of Harlaniella. Notice irregular development of
segmentation. Scale bar is 2 mm.
gues for a non trace fossil origin for all of these ‘‘ich-
nogenera.’’
Harlaniella and other cork-screw shaped forms
From several Neoproterozoic sections, rope-like
twisted worm-shaped objects have been reported. This
structure, known as Harlaniella Sokolov, 1972, is in-
terpreted by most specialists as a horizontal spiral trace
fossil, somewhat comparable to such Phanerozoic ich-
notaxa as Helicolithus and Helicorhaphe (e.g., Müller,
1971). In the basal Cambrian stratotype section at For-
tune Head, southeastern Newfoundland, Harlaniella
podolica is the nominative taxon for a Neoproterozoic
trace fossil Zone (Narbonne et al., 1987). Other oc-
currences of Harlaniella podolica include Podolia,
Ukraine (Kiryanov, 1968; Palij, 1976; Palij et al.,
1979); Lublin, and eastern Poland (Pacześna, 1986).
The spiral nature of Harlaniella appears to be based
on inferences of the three dimensional structure from
the rope-like surface morphology. However, Palij
(1976, p. 73) noted that Harlaniella podolica is similar
to Palaeopascichnus delicatus, and further (Palij,
1976, p. 74) that in Podolia, the two forms often are
found next to each other and may be transitional. A
similar relation appears to exist also in Newfoundland,
where Palaeopascichnus and Harlaniella occur on the
same bedding surfaces (Narbonne et al., 1987, fig.
6B). Palij’s significant observation appears not to have
SÖREN JENSEN
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been investigated further, and Harlaniella appears to
be accepted as a spirally twisted trace fossil.
The trace fossil interpretation of Harlaniella, may
need further consideration. This point may be illus-
trated with a specimen of Harlaniella from the Neo-
proterozoic of the Pasha core, south-east of lake La-
doga, western Russia (Fig. 6A, C), which was previ-
ously figured and described as Harlaniella sp. by Palij
et al. (1979; pl. 63:3). This specimen is similar to oc-
currences of Harlaniella podolica from Podolia illus-
trated by Palij et al. (1979, pl. 50:1–3). In two regions,
this specimen has inclined segments typical of Har-
laniella, but between these are transverse segments
identical to those seen in Palaeopascichnus (Fig. 6C).
In one of the outer regions, the inclined segments turn
sharply into an orientation nearly parallel to the struc-
ture. Overall, it is difficult to reconcile these obser-
vations with a spiral burrow.
A symmetrical spiral will follow a predictable
course and so fossil examples can be tested against the
expected model (Fig. 7). Application of this test to
specimens of Harlaniella shows that there are prob-
lems with the interpretation of Harlaniella as a helical
spiral. A specimen figured by Palij et al. (1979, pl. 50:
3) has in its central part segments that are imbricated
at about 458. The width of the trace is about 1.1 mm.
Assuming the extreme case that the width of spirally
coiled burrow is 0.55 mm, the distance between suc-
 PROTEROZOIC TRACE FOSSILS
FIG. 7. A. Cartoon depicting variable morphology of a spiral bur-
row related to angle of inclination to axis of trace. B. Tracing of a
portion of the holotype of Harlaniella podolica (taken from Sokolov,
1972). Case 1 and 2 depict the two examples of expected burrow
morphology if it was to meet theoretical properties of a spiral. Notice
that also in the most extreme case the spiral does not match the
morphology of the holotype.
cessive turns in a symmetrical spiral will be about 1.73
mm. The distance between segments on the fossil
range from 0.4–0.7 mm. Considering that the actual
width of the burrow in the fossil probably was less
than 0.55, it becomes even more difficult to make this
a spiral. Measurements for the type material of Har-
laniella figured by Sokolov (1972, 1973), yields sim-
ilar or larger discrepancies between actual and pre-
dicted measurements (Fig. 7).
Although compaction can be invoked, it would not
result in a consistent change in angle. It must therefore
be considered that Harlaniella is not a trace fossil. Its
close association with Palaeopascichnus (see above)
lends weight to this notion.
Large, apparently cork-screwed forms from South
Australia (Form A of Glaessner, 1969, fig. 5A) are
probably body fossils rather than trace fossils (Fedon-
kin and Runnegar, 1992).
Tubular body fossils
Preservation of tubular fossils as casts and molds on
clastic bedding planes appear to have been locally
common in the Neoproterozoic. These are easily con-
fused with trace fossils. Glaessner (1977) discussed
this problem in relation to whether Archaeichnium is
a trace or body fossil, but this problem appears to be
more extensive than has been realized previously. This
includes forms reported as Taenidum and Muensteria
from Namibia (Germs, 1972, pl. 2:2–3) which are
casts of a tubular fossil identical or similar to Clou-
dina. Other examples of probable tubular body fossils
are forms from South Australia that Jenkins (1995)
225
identified as Taenidum isp., and Palaeophycus tubu-
laris.
The difficulties involved in distinguishing trace fos-
sils from elongate tubular body fossils can be illus-
trated by a specimen that has been previously identi-
fied as Planolites (Palij et al., 1979, pl. 63:3). This
form shows a very gradual change in width which can-
not be explained solely by a trace changing level (Fig.
7A). In addition, parts of the structure show complex
internal fractures that are more consistent with fracture
of a brittle body wall than breakage of trace fossil fill
or lining (Fig. 7B). This structure is more readily ex-
plained as an elongate, gently tapering body fossil,
than as a trace fossil.
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In the examination of latest Neoproterozoic tubular
structures preserved as casts or molds in siliciclastic
sediment, more attention should probably be directed
to the increasing diversity of tubular organisms recov-
ered from the Doushantou Formation and other pre-
Ediacaran biotas (e.g., Xiao et al., 2002).
DISCUSSION
The view of Neoproterozoic trace fossils presented
above removes some of the peculiarities of certain oth-
er schemes. With the re-interpretation of forms such
as Palaeopascichnus there is no ‘‘extinction’’ of ich-
notaxa at the end of the Proterozoic. This also leads
to a decrease in the number of Neoproterozoic ich-
notaxa. Furthermore, it is possible to relate Neopro-
terozoic trace fossils more easily to Cambrian forms,
thus strengthening the case that they were produced by
metazoans. Arguable some of the more simple Neo-
proterozoic trace fossils could have had non-metazoan
producers (cf., Conway Morris, 1998), but this cannot
be the case for the more complex forms. At this point
must be considered the problems involved in reading
behavioral complexity from the morphology of trace
fossils. There exist no simple extrapolation of behavior
from trace fossil morphology, and even less a corre-
lation to neurological sophistication of trace producer.
It is, however, reasonable to argue that a complex trace
fossil represent a more complex behavior than does a
simple one. Miller (1998) suggested the useful terms
‘‘incidental’’ and ‘‘deliberate,’’ where incidental de-
note a simple structure likely of one function and short
occupation, whereas a deliberate trace fossils has a
more complex architecture likely representing multiple
functions and an extended time of occupation. One
type of complex trace fossils are those that consist of
numerous interconnected parts, of which Treptichnus
is a relatively simple example. Closely guided regular
meander traces is another type of trace fossil that re-
quires a moderately complex sensory systems (e.g.,
Seilacher, 1967). The presence of trace fossils such as
Spirorhaphe occurring with diverse Ediacaran forms
suggests that bilaterian evolution was well under way
at this stage. The Neoproterozoic trace fossils repre-
sent the initiation of a rapid but gradual build-up of
infaunal activity, which increased markedly in the
Cambrian.
226 SÖREN JENSEN
Problems in precise correlation of sections means
that only tentative suggestions can be made regarding
the evolution of Neoproterozoic trace fossils. A pre-
liminary attempt at such a correlation was presented
by Budd and Jensen (2000; fig. 6).
Narbonne et al. (1987) erected the Harlaniella po-
dolica Zone, corresponding to the upper part of the
Neoproterozoic trace fossil zone 1 of Crimes (1987).
This was defined by the occurrence of Palaeopasci-
chnus and Harlaniella. As discussed above these prob-
ably are not trace fossils.
As previously discussed (Jensen et al., 2000), trep-
tichnid trace fossils range below Treptichnus pedum in
Namibia and Spain. Treptichnids recently were found
below the first Treptichnus pedum in the Newfound-
land GSSP, though the precise implications of this re-
main unclear (Gehling et al., 2001). Treptichnids over-
lap the range of Cloudina in both Namibia and Spain
and thus far treptichnids have not been found in lo-
calities yielding diverse Ediacaran assemblages. The
treptichnids in Namibia are bracketed by radiometric
ages of 545 Ma and 548 Ma (Grotzinger et al., 1995)
and so are very close to currently accepted age for the
base of the Cambrian (e.g., Grotzinger et al., 1995).
Other trace fossils that first appear at approximately
this level are trace fossils with a three-lobed lower
surface (see Jensen and Grant, 1998). This suggest a
latest Neoproterozic trace fossil zone that may be cor-
relative with a low diversity Nama association (Gehl-
ing and Narbonne, 2002).
Sections with diverse Ediacaran body fossils are
dominated by simple horizontal forms and by the ap-
pearance of Archaeonassa. Radiometric dating of sec-
tions in the White Sea area suggest that these are youn-
ger than about 555 Ma (Martin et al., 2000). There are
reports of simple trace fossils of Planolites type that
are considered by some researchers to be older than
the main Ediacaran assemblages in Australia (e.g.,
Walter et al., 1989; Jenkins et al., 1992). This corre-
lation requires further confirmation but may suggest
an even older zone with short simple trace fossils.
Tentatively it is therefore possible to recognize two,
possibly three, latest Neoproterozoic trace fossil zones
(Fig. 8). New finds of trace fossils and further re-eval-
uations of old material coupled with rapidly increasing
new radiometric dates and revised correlations will test
the validity of this scheme. It is probably safe to argue
that trace fossils will continue to provide essential in-
formation on the early evolution of animals and that
it will ultimately be possible to devise a more detailed
and reliable Proterozoic zonation.
ACKNOWLEDGMENTS
My sincere thanks to Graham Budd and Kevin Pe-
terson for inviting me to this SICB Symposium, which
ultimately triggered me to publish ideas that had been
in manuscript form for years. For generously sharing
their material, time in the field, and stimulating dis-
cussions my sincere thanks to Graham Budd, Simon
Conway Morris, Mary Droser, Mikhail Fedonkin,
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FIG. 8. Trace fossil based zones at the Precambrian-Cambrian
boundary with brief characteristics. Lower Cambrian zones adapted
from MacNaughton and Narbonne (1999). The tentative Neoproter-
ozoic zones are proposed here (see text for discussion).
James Gehling, Kaisa Mens and Bruce Runnegar. The
majority of the research on which this paper is based
was conducted when I was at the Department of Earth
Sciences, University of Cambridge. I gratefully ac-
knowledge funding from the Leverhulme Trust and
NERC (grant GR3/10713 to Simon Conway Morris).
During my stay at Riverside I have benefited from
grants from National Geographic and the National Sci-
ence Foundation (grant EAR-0074021 to Mary Dro-
ser).
Robert Gaines read an earlier draft and significantly
improved the readability and language of this contri-
bution. James Hagadorn and an anonymous reviewer
are thanked for their insightful reviews.
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