Journal of the Geological Society, London, Vol. 165, 2008, pp. 307–318. Printed in Great Britain.

Sedimentation of the Phyllopod Bed within the Cambrian Burgess Shale Formation
of British Columbia

S . E . G A B B OT T 1, J. Z A L A S I E W I C Z 1 & D. C O L L I N S 2
1
Department of Geology, University of Leicester, Leicester LE1 7RH, UK (e-mail: sg21@le.ac.uk)
2
26 Belvedere Blvd, Toronto, Ontario, M8X1K1, Canada

Abstract: We provide the most detailed sedimentological log to date through the Phyllopod Bed of the mid-
Cambrian Burgess Shale Formation of British Columbia, based on millimetre-scale logging of a suite of thin
sections. The sedimentary facies is dominated by alternations of homogeneous mudstone and a coarser-
grained, laminated, variably sandy and shelly mudstone that is locally micronodular. Most boundaries between
these two lithologies are gradational, and discrete fining-upwards turbidite units were rarely recognized. Such
a pattern is interpreted to indicate rapid sedimentation of up to decimetre-thick units at this location from
pulsatory, quasi-continuous density currents consistent with earlier proposals of exceptional preservation
through rapid burial; the density currents responsible were probably largely akin to mud-rich slurries, helping
explain the transport and entombment of the fossils. The homogeneous mudstone units are characterized by
numerous distinctive lenses of pyrite framboids or subeuhedral crystals, previously interpreted as small
ripples. Their 3D shape, however, suggests an origin as subspherical early diagenetic aggregates; their present
morphology is consistent with the high levels of compaction inferred from the preservation of fossils.

The Burgess Shale is arguably the most celebrated fossil-bearing oxidant flux, to the extent that microbial decomposition was
unit in the world. Since its discovery by Walcott in 1909 and severely restrained. As well as restricted bioturbation owing to
particularly since the detailed re-descriptions of the faunas in the near-bottom water anoxia, low permeability was thought to be
1980s it has served as a key example of both the preservation of effected through deflocculated clays, early precipitation of pore-
soft-bodied faunas and the diversity of life soon after the occluding carbonate cements and an absence of coarse grains
Cambrian evolutionary ‘explosion’. Non-mineralized Burgess such as silts, microfossils, bioclasts and faecal pellets (Gaines et
Shale fossil remains are principally composed of kerogenized al. 2005).
films (Butterfield 1990), sometimes with associated aluminosili-
cates that according to Orr et al. (1998) replicated decay-prone
tissues prior to decomposition.
The sedimentology of the Burgess Shale has received rela-
tively little attention, despite the obvious significance that this
aspect has for the processes that lead to exceptional preservation.
Studies to date have interpreted the deposits as turbidites (e.g.
Piper 1972), and these have been used to invoke rapid burial
(obrution) of the fossils as a preservational mechanism (e.g.
Piper 1972; Whittington 1975; Conway Morris 1986; Allison &
Brett 1995). However, as turbidites are among the most common
of sedimentary facies worldwide, whereas Lagerstätten are by
definition rare, other factors to explain the exceptional preserva-
tion have been suggested. More recently, these have included
hypotheses to explain why Burgess Shale-type preservation (i.e.
kerogenized organic remains in a siliciclastic sediment; see
Butterfield 1990) has not been reported as a major taphonomic
pathway after the Cambrian. One suggestion relies on a pre-
ponderance of reactive and/or swelling clays in the sediments of
the Cambrian Gondwanan continental margins; these clays were
held responsible for prohibiting bacterial degradation of arthro-
pod cuticle and other reasonably recalcitrant organic tissues
(Butterfield 1995). Others (e.g. Allison & Briggs 1991, 1993; Fig. 1. Schematic representation of the stratigraphic relationships
Orr et al. 2003) have suggested that after the Cambrian the between the platform deposits of the Cathedral Limestone and Stephen
increase in the amount and complexity of bioturbation eliminated Formations and the basinal deposits of the Burgess Shale Formation (data
the deep-water low-oxygen taphonomic window, where charac- compiled from Fletcher & Collins 1998). The Burgess Shale Formation
teristically Burgess Shale-type preservation is found. Gaines et contains 10 members including the Walcott Quarry Shale Member, which
al. (2005) accounted for Burgess Shale-type preservation in the comprises the Greater Phyllopod Bed and the interval studied herein, the
Middle Cambrian Wheeler Formation of Utah through a combi- Phyllopod Bed. WLM, Wash Limestone Member; GB, ‘Ginger Bed’;
nation of influences that reduced sediment permeability, and thus GML, ‘Great Marrella layer’; GEL, ‘Great Eldonia layer’.

307
308 S . E . G A B B OT T E T A L .

To shed more light on the processes involved in the deposition
of the Burgess Shale we have sampled a large proportion of the
Phyllopod Bed from one vertical section at the southern side of
Walcott’s Quarry. The sedimentary facies were logged from thin
sections at a millimetre scale. The Phyllopod Bed is the best-
known and most fossiliferous unit of the Walcott Quarry Member
(Burgess Shale Formation). The results of our analyses provide
significant constraints on the depositional processes involved,
and allow insights into the mechanisms of fossil preservation at
this locality.
 S E D I M E N TAT I O N O F T H E P H Y L L O P O D B E D
Previous sedimentological studies
The Phyllopod Bed has been described as comprising sharp-
based units of calcareous siltstone grading up via interlamination
into mudstone by Piper (1972), who interpreted these units to be
309
Fig. 2. Graphic log alongside composite
image of the thin sections of the Phyllopod
Bed. Heights are in centimetres starting at
0 cm, which is the base of the Phyllopod
Bed just above the ‘Ginger Bed’. White
chevrons on the thin-section image indicate
positions where lateral movement along a
lamina was necessary to capture the
complete image; it should be noted that in
these cases there is no stratigraphic break.
Grey arrows show positions of sharp
boundaries. Stratigraphic gap represents an
interval where no rock was collected or an
interval where the rock splintered so that
thin sections could not be made. ‘Sliver
missing’ indicates position where a
maximum of 2–3 mm is missing as the
shale splintered into small pieces unsuitable
for thin sectioning. The left-hand side of
the graphic log indicates the presence of a
diagenetic carbonate cement and the right-
hand side of the graphic log indicates
features such as pyrite lenses, shell
fragments and bedding structures.
turbidites, where the calcareous siltstone was derived from the
nearby Cathedral Formation reef; the muddy deposits and fossils
were interpreted as eroded by the turbidity currents from
intermediate depths. Piper (1972) described a typical turbidite
unit as comprising, at its base, a lower calcareous siltstone with
310 S . E . G A B B OT T E T A L .

Fig. 3. Photographs to demonstrate the nature of facies seen in the Phyllopod Bed. Heights provided indicate height above the Ginger Bed for the base of
each photograph shown. (a) The homogeneous mudstone facies from the Great Marrella layer. White arrows and the black arrow indicate pyrite lenses
and a ‘floating’ quartz grain, respectively. Height 1 cm; scale bar represents 2 mm. (b) Coarser poorly sorted facies: pale mottled layers represent varying
degrees of diagenetic carbonate cement. Height 82.2 cm; scale bar represents 2 mm. (c) Image showing the different textural fabrics (representing different
degrees of cementation) within the coarser poorly sorted facies. Height 85.2 cm; scale bar represents 2 mm. (d) BSE image showing a coarser poorly
sorted layer with carbonate cement in the centre and its gradational boundaries with the adjacent homogeneous mudstone. Height 86 cm; scale bar
represents 500 ìm.

distinctly irregular laminae (mudstone laminae being absent), thickness of mudstone laminae; upwards in each unit, mudstone
overlain by alternating laminae of mudstone and well-packed laminae were described as alternating with carbon-rich laminae
calcareous siltstone, with a decrease upwards in the thickness, including some calcareous silt-sized clasts, this grading into the
coarseness and packing of the siltstones, and an increase in the highest part of the unit, comprising mudstone with a small
 S E D I M E N TAT I O N O F T H E P H Y L L O P O D B E D
proportion of carbonate and no visible laminae. Piper (1972)
suggested that many of the unlaminated mudstone units towards
the upper portion of the Phyllopod Bed were hemipelagic. Our
interpretation of the basic sedimentological pattern (see below)
differs substantially from this.
Allison & Brett (1995) have also reported details of deposits at
Walcott’s Quarry. They described intercalations comprising mas-
sive thicker beds and finely laminated thinner beds. The laminae
were commonly seen to fine upwards, to have erosive bases with
small microscours, 2–3 mm across, and some lamina bases
contained detrital quartz, carbonate fragments, and occasional
rip-up clasts. They identified small ripples, some formed from
pyrite framboids (Allison & Brett 1995, fig. 2c, p. 1080), and
reported that in deposits from Walcott’s Quarry pyrite is almost
invariably evenly distributed, with rare clustering, except for thin
pyrite patinas associated with worm gut traces. Allison & Brett
(1995) proposed that organisms preserved in the Phyllopod Bed
were engulfed in high-density mud–silt flows; an interpretation
consistent with our conclusions. Gostlin & Miall (2005) reported
calcisiltite layers intercalated with massive, sharp-based clay-rich
mudstones from the Greater Phyllopod Bed. They stated that the
presence of massive beds and high clay contents were incon-
sistent with deposition of the Burgess Shale via turbidity currents
and fluidized mudflows, and suggested that deposition of muddy
sediments and fauna occurred through settling after storm-gener-
ated back-currents swept sediment into the basin.
Sampling and methods
The section targeted was the whole of Walcott’s original Phyllopod Bed
(Walcott 1912), which is just over 2 m thick at this site. The Phyllopod Bed
begins immediately above the informally designated ‘Ginger Bed’, which
is an ochreous, pyritic arenaceous bed, and extends to the top separation
plane of Walcott’s Quarry (Fletcher & Collins 1998). The Phyllopod Bed
constitutes part of the Walcott Quarry Member of the Burgess Shale
Formation (see Fletcher & Collins 1998); it is the most prolific of
fossiliferous beds within this formation (Whittington 1985; Conway Morris
1986), containing the classic Burgess Shale fauna, and was the principal
focus of previous studies by Piper (1972) and Allison & Brett (1995). The
latter also studied material from Raymond’s Quarry, which lies c. 35 m
higher up-section (Allison & Brett 1995). The Phyllopod Bed was
discovered by Walcott in 1909, and was further excavated by the
Geological Survey of Canada (1966–1967). Subsequent excavations (from
1993 to 2000) led by one of us (D.C.) extended 5 m down from Walcott’s
original quarry floor (and coincident base of the Phyllopod Bed) to the top
of the Wash Limestone. Thus the ‘Greater Phyllopod Bed’ is a stratigraphic
interval extending from the top of the Wash Limestone to the top of the
Phyllopod Bed and is about 7 m thick (Fig. 1).
The Phyllopod Bed has been logged in considerable detail in the field
by Fletcher & Collins (1998), who recognized significant lateral variation
in bed thicknesses. One of us (S.G.) joined the Royal Ontario Museum
field crew and collected samples from the Walcott’s Quarry Member,
including a sequence of samples from the Phyllopod Bed. The log of
Fletcher & Collins (1998, fig. 4, p. 419) provided a framework in which
the Ginger Bed comprises a marker bed that lies directly below the
Phyllopod Bed, and heights are measured from 0 cm, which defines the
base of the Phyllopod Bed (and the top of the Ginger Bed). The samples
collected from the Phyllopod Bed cover 60% of the total interval; there
are only four significant gaps between 12 and 42 cm (30 cm missing), 59
and 68 cm (9 cm missing), 70 and 78 cm (8 cm missing) and 168 and
186 cm (18 cm missing); here the rock splintered into small pieces as
collection was attempted. Despite this, the material collected represents
the most complete set of samples for detailed lamina-scale sedimentolo-
gical analysis of the Burgess Shale yet collected. Two highly fossiliferous
layers were reported by Walcott (1912) and constitute useful marker
horizons in the Phyllopod Bed; they are the ‘Great Marrella layer’ (0–
311
7.4 cm), which clearly identifies the base of the section, and the ‘Great
Eldonia layer’ (125–130 cm).
A suite of polished thin sections of the entire rock succession collected
was prepared. The thin sections were logged at a millimetre scale using a
PetroScope1 to create the log shown in Figure 2. After carbon coating,
backscattered electron (BSE) imagery was obtained using a Hitachi
scanning electron microscope (S3600N). Elemental analyses of minerals
were determined by energy-dispersive spectrometry using an Oxford
Instruments INCA system. Photographs of the thin sections (Figs 3–5)
were obtained by placing the thin sections directly into a Durst M805
enlarger and exposing photosensitive paper.
Sedimentary facies
The sedimentary facies essentially comprises a continuum be-
tween two end-members.
(a) The first end-member is a fine-grained homogeneous
mudstone facies (Figs 3a and 4a–d) that originally was dom-
inantly composed of clay minerals (illite–smectite–kaolinite);
these were recrystallized to muscovite–chlorite–quartz–albite
during greenschist-grade metamorphism (Powell 2003). This
facies includes a minor component of silt-sized quartz grains
that, in general, become more numerous as the homogeneous
mudstone facies grades into the coarser end-member described
below (Fig. 4c and d); some larger (sand-sized) matrix-supported
(‘floating’) quartz grains and shell fragments are also locally
present (Figs 3a and 4a–d). A feature of this facies is the
occurrence of lenses (up to 1000 ìm, but more commonly 200–
500 ìm in length) composed of pyrite framboids and more rarely
pyrite euhedra (Figs 3a, 4a,c and 6). Intervals over 7 cm thick of
essentially massive, ungraded mud occur; these include the
‘Great Marrella layer’ (Fig. 2 (0–7.4 cm) and Fig. 3a) and the
‘Great Eldonia layer’ (Fig. 2 (125–130 cm)).
(b) The second end-member is a coarser facies characterized
by poorly sorted, subordinate quartz silt and sand grains (up to
1.5 mm in diameter, but usually 150–500 ìm in diameter) with
shelly fossil fragments up to 4 mm in size within a mud matrix
(Figs 3b–d, 4e and 5a,d). The larger particles, both of quartz and
detrital carbonate, conspicuously ‘float’ within a finer mud
matrix (Figs 4e and 5a,d). Where present, a locally abundant
diagenetic carbonate component overprints most of the primary
lamination (Fig. 3d). Typically, this facies shows a mottled
texture with pale carbonate lensoids surrounded by darker
mudstone (usually the coarser end-member), but the carbonate
lensoids also locally coalesce to give more massive carbonate-
cemented layers between which there are bedding-subparallel,
wispy, muddy intercalations (e.g. Figs 3b,d and 4e). The mottled
textures are, in places, accentuated by stylolitic development (see
Powell 2003). In the coarser, poorly sorted facies, pyrite is
irregularly disseminated or forms bedding-parallel wisps, and
only very rarely occurs as the discrete lenses typical of the
homogeneous mudstone facies.
These end-members, (a) and (b), intergrade in vertical succes-
sion in both fining-upwards (e.g. Fig. 2) and coarsening-upwards
(e.g. Figs 2 and 5b) units; broadly centimetre-scale trends are
common (Fig. 2). Coarsening-upwards trends are preponderant
(35 being identified, compared with 24 fining-upwards trends)
suggesting the frequent incidence of waxing flow events. Clear
interfaces that may represent breaks in sedimentation are rare
(only eight were identified; see Figs 2 and 4) and, where present,
they are commonly relatively fine-based. These might represent
time gaps involving cessation of deposition; we cannot constrain
the duration of these gaps other than to note that neither
identifiable hemipelagic laminae nor bioturbated intervals are
associated with these interfaces. The intergradation between
312 S . E . G A B B OT T E T A L .
 S E D I M E N TAT I O N O F T H E P H Y L L O P O D B E D 313

coarser and finer mud ranges from centimetre scale to millimetre
scale. Higher in the succession (starting at 109 cm), and
associated with a general increase in the proportion of coarser
material, there are local intervals of inclined millimetre-scale
laminae up to 2 mm thick (Fig. 5c) that may represent ripple
cross-lamination. We found no textures that could be clearly
associated with bioturbation.
Pyrite lenses
A characteristic feature of the homogeneous mudstone units
within the Burgess Shale are small (200–800 ìm in length)
bedding-parallel lenses, each composed of some dozens to
hundreds of pyrite framboids or, less commonly, of euhedral to
subhedral crystals (see Figs 3a, 4a and 6). These have been
previously observed and interpreted as small-scale ripples of
transported pyrite grains (Allison & Brett 1995, fig. 2C, p. 1080).
We have observed these on bedding surfaces, where they have
a roughly circular distribution (Fig. 6c and f); consequently, their
overall geometry is that of ovoids that are highly shortened along
the vertical axis; this is not consistent with ripple-forms. As far
as we could judge, these pyrite lenses showed no spatial
association with fossils or organic fragments, although patinas of
pyrite are locally associated with both soft tissue and skeletal
fragments in the Burgess Shale (e.g. Whittington 1975; Conway
Morris 1985, 1986; Allison & Brett 1995).
The textures seen by BSE imagery are generally consistent with
a diagenetic origin of the framboids in the lenses, and the large size
of most of the framboids (e.g. mean diameter for framboids in the
pyrite lens shown in Fig. 6a is 9.7 ìm) suggests precipitation
within the sediment rather than within the seawater column (see
Wignall & Newton 1998). Powell et al. (2003) suggested that pyrite
framboids in the Burgess Shale displayed significant evidence of
recrystallization and accordingly that the statistical analysis of
framboid size as a palaeoredox indicator was limited. However, the
framboids measured here from the lens in Figure 6a do not show
the recrystallization features reported by Powell et al. (2003), such
as solid spheroidal grains of similar size to framboids, and so we
interpret them as original. The pyritic lenses are strongly bedding-
parallel even in mudstone that otherwise appears perfectly homo-
geneous, and so growth of the framboids along some pre-existing
bedding-parallel fabric can be precluded. Thus, in attempting an
interpretation, both the clustering of the framboids into the lenses
and the alignment of the lenses need to be addressed.
Tentatively, we link the clustering of the framboids into the
lenses with the rapid sedimentation we infer for the entire unit
(see below). Thus, we envisage that the sudden burial of a mass
of sediment, initially containing a high content of (at least partly
oxygenated) entrapped seawater, might produce, fleetingly (be-
fore significant compactional dewatering began), a broad rela-
tively permeable zone of redox contrasts (i.e. between sediment
particles and seawater) that led first to initial ‘random’ precipita-
tion of framboids. Subsequently, we suggest that rapid diffusion
of iron and sulphide ions to local centres of precipitation would
have taken place, the diffusion paths being driven by concentra-
tion gradients produced by the pyrite crystallization process itself
(Fig. 7); roughly spherical aggregates of pyrite framboids would
result. Subsequent application of the considerable amount of
compaction (a minimum compaction ratio for the Burgess Shale
of 8:1 was inferred from fossils by Whittington (1975)) would
result in the transformation of the spherical aggregates of
framboids into highly flattened ellipses. Thus, their present shape
and alignment, in this interpretation, reflects compaction directly,
and bedding only indirectly.
Interpretation and discussion
The pattern of lamination observed does not accord with the
previously published sedimentological description of Walcott’s
Phyllopod Bed as a succession of rhythmic couplets of a simple
turbidite model where each couplet represents sedimentation
from a discrete turbidity current (Piper 1972). Rather, we have
observed a considerably less ordered pattern that shows a
succession of gradations between relatively coarser and relatively
finer sediment with reverse graded intervals slightly more
common than normally graded intervals. Reverse grading and
‘floating’ outsized clasts and bioclasts in an otherwise ungraded
mud (facies (a) described above) seems also not consistent with
the model proposed by Gostlin & Miall (2005) of settling of
material from the water column after storms.
Likewise, the facies pattern we have observed seems not
consistent with deposition either by hemipelagic processes or
from more or less continuous sea-floor currents (i.e. as contour-
ites). Modern hemipelagites are mostly intensely bioturbated and
so do not provide a good comparison. Better comparison is made
with hemipelagites described from early Palaeozoic basins; for
example, those from the central Welsh basin (Cave 1979; Davies
et al. 1997). Here, hemipelagites that accumulated on an
essentially anoxic sea floor, as were prevalent in those times,
show a clearly laminated structure with organic-rich (pelagic)
laminae alternating with clastic laminae deposited from nephe-
loid plumes. At intervals when the sea floor was oxygenated this
lamina structure was visibly disrupted by bioturbation. None of
the deposits we describe resemble this widespread early Palaeo-
zoic facies. Similarly, although our idealized log (Fig. 8) super-
ficially resembles the idealized contourite of Stow et al. (2002:
p. 18, fig. 10) the gradational boundaries of the latter are
achieved through pervasive bioturbation, a phenomenon that we
have not observed in our material from the Phyllopod Bed.
The pattern observed, with only eight boundaries where a
significant break in deposition may be inferred in the material we
have studied (over 60% of the Phyllopod Bed), is more consistent

Fig. 4. Sharp boundaries in the Phyllopod Bed. Heights provided indicate height above the Ginger Bed for the base of each photograph shown. (a)
Homogeneous mudstone facies with three conspicuous pyrite lenses (white arrows) and a shell fragment (black arrow) sharply overlying less well-sorted,
coarser mudstone facies; this is the sharpest, most distinct boundary within the sampled Phyllopod Bed. Height 57.7 cm; scale bar represents 2 mm. (b)
Relatively sharp boundary lying just above the large carbonate clast (note large bright quartz clast just to right of this) towards bottom of image, between
finer (below) and coarser (above) mudstone layers. Upper half of image shows gradational contacts between finer and coarser layers, both of which
contain ‘floating’ clasts (e.g. shell fragment labelled with white arrow). Height 7.4 cm just above the ‘Great Marrella layer’; scale bar represents 2 mm.
(c) Rapidly gradational boundaries between coarser (with abundant silt and fine sand) and finer mudstone layers. (Note pyrite lenses (white arrows) in
finer mudstone layer and large detrital quartz clast (white) in the coarser layer.) Height 68.5 cm; scale bar represents 2 mm. (d) BSE image of moderately
sharp boundary between coarser mudstone with a little carbonate cement (below) and finer mudstone layers. Poorly developed pyrite lens (white arrow)
and ‘floating’ quartz clast (black arrow). Height 79 cm; scale bar represents 500 ìm. (e) Image showing the variable nature of boundaries between layers
of different grain size from sharp (white arrow) and irregular gradations (bottom and top parts of image). (Note the ‘floating’ shell debris.) Height
143.8 cm; scale bar represents 2 mm.
314 S . E . G A B B OT T E T A L .
 S E D I M E N TAT I O N O F T H E P H Y L L O P O D B E D 315

Fig. 6. BSE images of pyrite lenses. (a) Cross-section through a pyrite lens in the Great Marrella layer composed of framboids of varying sizes. Scale bar
represents 100 ìm. (b) Close-up of pyrite lens shown in (a): the framboids are moderately disordered but show no evidence of overgrowth or alteration;
microcrystals are euhedral to subhedral. Scale bar represents 50 ìm. (c) Bedding-parallel image to show a pyrite lens in the Great Marrella layer. (Note
that the framboids have a roughly circular distribution parallel to bedding.) Scale bar represents 100 ìm. (d) Cross-section through a pyrite lens where the
pyrite crystals are more closely packed than framboids forming the lens in (a) and (b). Height c. 89.5 cm; scale bar represents 100 ìm. (e) Close-up of the
pyrite lens shown in (d), showing pyrite crystals ranging from anhedral to euhedral (octahedral and cubic) in habit. Scale bar represents 50 ìm. (f)
Bedding-parallel image to show a pyrite lens in the Great Eldonia layer. (Note the circular distribution of the octahedral pyrite crystals.) Scale bar
represents 30 ìm. (g) Framboids in pyrite lens from the Great Eldonia layer where the microcrystals are fairly disordered: these are moderately tightly
packed in the largest framboid but are loosely packed in the other framboids; microcrystals are cubo-octahedral. Scale bar represents 10 ìm.

with pulsatory deposition from waxing and waning semi-contin-
uous density currents producing successive packets of rapidly
accumulated sediment (see Best et al. 2005). Some of these units
would have been at least decimetres thick prior to compaction;
for example, the Great Marrella Bed (here 7 cm thick post-
compaction).
In terms of the Bouma model the single intervals most closely
compare with units Td (the millimetre-scale coarse–fine inter-
gradations) and Te (the homogeneous mudstones) with frequent
gradations between these. Locally, near the top of the section
studied, there are thin intervals consistent with the rippled Tc
unit. However, such comparisons with the Bouma model (or with

Fig. 5. Gradational boundaries in the Phyllopod Bed. Heights provided indicate height above the Ginger Bed for the base of each photograph shown. (a)
Typical section through the upper part of the Phyllopod Bed showing rapid gradational boundaries between units of different grain size and sorting, and
variable expression of the superimposed micronodular, carbonate cement fabric in the coarser layers from discrete bedding-parallel ‘laminae’ to irregular
mottling. Scattered pyrite lenses in finer layers (white arrows) and ‘floating’ quartz clasts and shell fragments throughout. Height 154 cm; scale bar
represents 2 mm. (b) Typical gradational coarsening-upwards trend in generally homogeneous silty mudstone. Height 47 cm; scale bar represents 1 mm.
(c) Overall fining-up succession with lenticular and wispy lamination (indistinct ripple forms?) in lower part of image. Height 109 cm; scale bar represents
2 mm. (d) Lower half of image shows silty mudstone with large ‘floating’ shell fragment (white arrow); this facies gives way upwards to alternating layers
differing in grain size and with variable degrees of micronodular, carbonate cement (pale). Height 165.3 cm; scale bar represents 2 mm.
316 S . E . G A B B OT T E T A L .
Fig. 7. Schematic illustration showing one possible model for the
formation of the lenses comprising pyrite framboids, and more rarely
euhedral to anhedral crystals. (a) Highly uncompacted sediment layer as
mud–seawater mixture following deposition from density currents; initial
randomly distributed precipitation of pyrite framboids. (b) Creation of
Fe2þ and sulphide ion species concentration or diffusion gradients
favouring further pyrite nucleation in localized, roughly spherical
volumes of sediment. (c) Compaction of the sediment by .85% to create
pyrite framboid clusters as flattened ovoids (‘pancake shapes’).
Fig. 8. Schematic logs comparing the distribution of clastic particles
(including fossils) between a typical early Palaeozoic turbidite facies,
such as the early Silurian strata of central Wales, (a) and the Phyllopod
Bed (b). It should be noted that the mudstone layers in the Phyllopod
Bed contain outsized clasts (including fossils), whereas these are absent
from the turbidite mudstone layers in the Welsh example depicted.
Hemipelagite facies have not been recognized by us in the Phyllopod
Bed, but are an integral and distinctive component of the Welsh
turbidites, either as laminated organic-rich layers laid down in anoxic
sea-floor conditions (AHP) or penecontemporaneously oxidized and
burrowed layers (OHP), if laid down on an oxygenated sea floor.
the finer subdivisions of the Td and Te units proposed by Piper
(1978) and Stow (1979)) are probably an oversimplification, for,
characteristically in the Phyllopod Bed, over-sized quartz grains
and shell fragments commonly ‘float’ within finer-grained sedi-
ment accompanying the macrofaunal animal remains. This
indicates that the density current possessed, at least over the
distance between the source of the large clasts and the aggrading
surface (see Branney & Kokelaar 2002), sufficient competence to
transport such material. This in turn implies deposition not from
dilute turbidity currents (by sedimentation of mud particles from
suspension), but from denser, mud-rich suspensions that, at least
intermittently, were perhaps akin to slurries, or ‘slurry-flows’
(i.e. flows transitional between turbidity currents and debris
flows: Lowe & Guy 2000; Lowe et al. 2003) (see also Mulder &
Alexander 2001; Amy et al. 2006). Given the frequency of both
reverse and normal grading in the Phyllopod Bed, we prefer
interpretations involving progressive aggradation of the waxing
and waning of such flows rather than models involving en-masse
freezing of high-density, non-turbulent currents (e.g. McCave &
Jones 1988).
For comparison, in mud-dominated turbidite deposits, such as
those of the Welsh Basin (Davies et al. 1997), that portion of the
turbidity currents responsible for the deposition of the Bouma D
and E intervals was typically insufficiently competent to transport
and entomb graptolite rhabdosomes (which would mostly have
been hydrodynamically lighter than most of the Burgess animals,
and thus more easily transported). Graptolites in these rocks are
typically found as current-sorted accumulations in rippled Bou-
ma C intervals (Davies et al. 1997; Zalasiewicz 2001) of
turbidite units, whereas overlying Bouma E mud layers are
relatively well-sorted, fine-grained and contain neither trans-
ported graptolite remains nor outsized mineral grains (Fig. 8).
This variation on the standard turbidite model, as regards the
Phyllopod Bed, may well be a factor in the exceptional fossil
preservation observed within this unit (see below). The best
preservation of non-mineralized fossils occurs in intervals of the
Burgess Shale that are within the finer-grained, more homoge-
neous units within our classification; for example, the ‘Great
Marrella layer’ and the ‘Great Eldonia layer’. Our interpretation
of the depositional process is consistent with the concept of rapid
burial (at least within the interval that we have studied) outlined
by earlier workers (e.g. Whittington 1975, 1980; Conway Morris
1986). In addition, our interpretation of these units as having
been deposited from relatively dense slurries (in which turbu-
lence may have been damped), rather than as typical Bouma E
units, is consistent with the size of the animals or carcasses being
transported (Fig. 8) and the lateral variability in bed thicknesses
recognized by Fletcher & Collins (1998).
Furthermore, the location of these units within a broader
interval (the Phyllopod Bed), which shows signs of having been
rapidly accumulated overall, strengthens the argument that rapid
burial is a key factor in exceptional preservation at this locality.
If the Phyllopod Bed was indeed deposited effectively as a small
number of units, from pulsed, dense, mud-rich slurries, and if the
compaction factor of .85% deduced from the entombed fossils
(Whittington 1975) and the geometry of the pyrite lenses (see
above) is broadly correct, then one may envisage a geologically
instantaneous (and thus ‘catastrophic’) accumulation of deci-
metre- to metre-scale thicknesses of sediment. The entombed
animals would be prevented from floating away as they accumu-
lated decay-generated gases and many would also thus be
‘instantly’ taken below the highly bacterially active surface layers
of sediment.
This scenario contrasts sharply with typical centimetre- or
decimetre-scale turbidites that, again drawing analogy with the
Welsh Basin, were separated by decadal or centennial intervals
during which slow hemipelagic sedimentation and sea-floor
chemical and biological activity took place (see Fig. 8; Cave
1979; Davies et al. 1997). All of the laminated intervals of the
 S E D I M E N TAT I O N O F T H E P H Y L L O P O D B E D
Phyllopod Bed have a texture generally consistent with tractional
lamination. We observed no interval that appeared to represent
hemipelagic deposition, as is typical, for instance, of the deep-
water graptolite shales of the Early Palaeozoic (e.g. compare
Armstrong & Coe 1997; Davies et al. 1997, plate 10A–C, p. 65).
Similarly, we have seen no evidence of microbial mats as
reported by Powell et al. (2003) and Caron & Jackson (2006),
although these come mostly from below the Phyllopod Bed. We
cannot preclude the existence of brief pauses to allow the growth
of microbial mats, or to allow brief intervals of colonization
(sporadic burrows have been noted in this interval; Fletcher &
Collins 1998), but we saw no evidence of these associated with
the sharp interfaces recognized in our material.
How do our data reflect upon the status of the Burgess Shale
fossils as ‘census’ life assemblages (i.e. a fossil assemblage
composed of species belonging to a single community and
preserved in the environment in which they lived), death
assemblages (i.e. a transported fossil assemblage), or time-
averaged assemblages (i.e. an accumulation of fossil species,
over a period of time)? The Great Marrella layer and the Great
Eldonia layer are striking examples of the homogeneous mud-
stone facies, with evidence of subtle waxing and waning flow,
and a lack of interfaces, that we deduce represents material
transported as substantial gravity-driven units akin to slurry
flows. Hence, the animals entirely enclosed in these layers were
subjected to some transport and were not buried in situ. It is
perhaps possible that some animals were able, if buried in situ,
to migrate up through the newly deposited layer. However, we
see no evidence of, for instance, escape traces, nor of fossils
associated (as colonizers) with the few sedimentary interfaces
that we have recognized. Moreover, the fauna in the Great
Marrella layer and Great Eldonia layer includes taxa (e.g.
Scenella, Selkirkia and algae: Caron & Jackson 2006) unlikely to
be capable of moving through tens-of-centimetres thickness of
suddenly deposited sediment.
However, we cannot constrain the distance of transport, which
may have been minimal, nor the relative coherence of the
assemblages as communities. Caron & Jackson (2006) have
demonstrated that single beds in the Great Phyllopod Bed
(including the Great Marrella layer and Great Eldonia layer)
contain articulated organisms, interpreted as census assemblages
and in situ dissociated and completely dissociated organisms,
interpreted as time-averaged assemblages. Their analyses of
more than 50 000 specimens indicated that, although many
organisms were moved, they were not transported out of their
community habitat, and hence the often cited concept of a pre-
slide environment (where the animals lived) and a post-slide
environment (where they ended up) was not valid (Caron &
Jackson 2006). Some degree of transport is consistent with
reported occurrences of numerous organisms preserved at
various angles with respect to bedding, the existence of
sediment between appendages and the preferred (current-
aligned) orientations of Selkirkia tubes (Conway Morris 1986).
However, we cannot preclude that some of the fossils in the
Phyllopod Bed (although not those in the Great Marrella layer
and Great Eldonia layer) may have been buried in situ; this is
suggested by the presence of trilobite and other arthropod
moults in the Great Phyllopod Bed (D.C., personal observation;
Caron & Jackson 2006).
The rate of deposition we infer, together with the absence of
recognizable hemipelagic deposits, suggests that it is difficult
to constrain the oxygenation state of the Phyllopod Bed (see
Cave 1979; Davies et al. 1997). The recognition of sporadic
burrowed intervals more broadly within the Burgess Shales
317
(e.g. Allison & Brett 1995; Powell et al. 2003; Caron &
Jackson 2006) suggests a sea floor that was at least intermit-
tently oxic, and the presence or absence of burrows has been
used to infer changes in basin oxicity (Caron & Jackson
2006). Our analysis suggests that the perceived absence of
burrows may here be due to inhibition of colonization through
rapid sediment accumulation, where bioturbators were unable
to keep pace with sediment influx, as much as by sea-floor
and/or sediment anoxia.
The pervasive occurrences of the distinctive pyrite lenses and
our inference of a diagenetic origin for them (see above) suggests
a distinctive geochemical environment of early diagenesis in the
Burgess Shale perhaps linked with the distinctive mode of
deposition we infer. The originally uncompacted state of the
deposits inferred from our results and from fossils that are found
at various angles to bedding (locally very high angles despite
subsequent compaction) suggests that the preservation model of
Gaines et al. (2005), for the Wheeler Formation Lagerstätte of
Utah, cannot be used to explain fossil preservation in the Burgess
Shale. Those workers showed that a number of factors resulted in
sediments with much reduced permeability, a situation further
compounded by early, ubiquitous pore-occluding, carbonate
cementation. Gaines et al. (2005) suggested that preservation of
kerogenized carbon films in the Wheeler Formation sediments
was the result of low permeability that lowered oxidant flux
sufficiently to restrict microbial decay. However, rapid sedimen-
tation from mud-rich density currents would have produced
relatively high initial levels of porosity and permeability in the
Burgess Shale. This is suggested both by estimates of the
compactional flattening of enclosed fossils (Whittington 1975)
and pyrite framboid aggregates (see above) and by the high
porosities of modern muddy sediments immediately after deposi-
tion (commonly up to 80% by volume: Singer & Muller 1983,
p. 188). Subsequent burial would squeeze out pore fluid, but
probably not on the short time scale necessary for effective
operation of a low-permeability preservational mechanism. In
addition, unlike the Wheeler Formation, the Burgess Shale of the
Phyllopod Bed does contain skeletal bioclasts and larger detrital
grains that would have served to increase primary porosity. Our
observations thus suggest burial rate rather than porosity as a
major factor.
High levels of compaction may have produced significant
diagenetic fissility in the Burgess Shale. This, with heating to
250–280 8C (I. Harding, pers. comm., in Butterfield 1995) upon
burial to 10 km (Powell 2003), could have produced the reported
S1 bedding-parallel ‘cleavage’ (Powell 2003) by mimetic recrys-
tallization on the fissility. As these rocks were not isoclinally
folded prior to the Mesozoic Laramide Orogeny, and as over-
burden (lithostatic) pressure itself does not produce a directional
fabric, this might represent a better explanation for the ‘cleavage’
reported from these rocks.
Our observations and inferences help provide an explanation
for the exceptional preservation in the Burgess Shale, but may
not offer much insight into the wider problem of the concentra-
tion of Lagerstätten around continental margin and shelf-basin
environments in the Cambrian (Allison & Briggs 1993). The
perceived high relative abundance of Cambrian Lagerstätten has
been linked to physico-chemical conditions, such as specific clay
chemistries (Butterfield 1995) and factors reducing permeability
(Gaines et al. 2005), or to a post-Cambrian increase in the
amount and complexity of bioturbation, which effectively elimi-
nated the deep-water slope-basin taphonomic setting (Allison &
Briggs 1991, 1993, 1994; Orr et al. 2003; but see Aronson 1992;
Pickerill 1994). Whatever the solution to this general problem,
318 S . E . G A B B OT T E T A L .
detailed sedimentological analysis can shed substantial light upon
local instances of exceptional preservation.
The fieldwork for this project could not have been done without the
agreement and support of the Canadian Parks Service. S.G. in particular
thanks fieldwork crew members J. B. Caron, D. Garcı́a-Bellido, K.
Gostlin and M. Myers. Financial help was provided by a University of
Leicester Research Support Grant to S.G. and a Royal Society Research
Equipment Grant to S.G. and J.Z. L. Barber drafted Figure 2. We thank
P. Allison and L. Amy for their incisive and helpful reviews, T. Fletcher
for valuable comments on an earlier draft, and M. Branney, J. Macquaker,
A. Page and J. Schieber for useful discussions.
References
Allison, P.A. & Brett, C.E. 1995. In situ benthos and paleo-oxygenation in the
Middle Cambrian Burgess Shale, British Columbia, Canada. Geology, 23,
1079–1082.
Allison, P.A. & Briggs, D.E.G. 1991. Taphonomy of nonmineralized tissues. In:
Allison, P.A. & Briggs, D.E.G. (eds) Taphonomy: Releasing the Data
Locked in the Fossil Record. Plenum, New York, 26–71.
Allison, P.A. & Briggs, D.E.G. 1993. Exceptional fossil record: distribution of
soft-tissue preservation through the Phanerozoic. Geology, 21, 527–530.
Allison, P.A. & Briggs, D.E.G. 1994. Exceptional fossil record: Distribution of
soft-tissue preservation through the Phanerozoic: Reply. Geology, 22, 184.
Amy, L.A., Talling, P.J., Edmonds, V.O., Sumner, E.J. & Lesueur, A. 2006. An
experimental investigation of sand–mud suspension settling behaviour;
implications for bimodal mud contents of submarine flow deposits. Sedimen-
tology, 53, 1411–1434.
Armstrong, H.A. & Coe, A.L. 1997. Deep-sea sediments record the geophysiol-
ogy of the Late Ordovician glaciation. Journal of the Geological Society,
London, 154, 929–934.
Aronson, R.B. 1992. Decline of the Burgess Shale fauna: ecologic or taphonomic
restriction? Lethaia, 25, 225–229.
Best, J.L., Kostaschuk, R.A., Peakall, J., Villard, P.V. & Franklin, M. 2005.
Whole flow field dynamics and velocity pulsing within natural sediment-laden
underflows. Geology, 33, 765–768.
Branney, M.J. & Kokelaar, P. (eds) 2002. Pyroclastic Density Currents and the
Sedimentation of Ignimbrites. Geological Society, London, Memoirs, 27.
Butterfield, N.J. 1990. Organic preservation of non-mineralizing organisms and
the taphonomy of the Burgess Shale. Paleobiology, 16, 272–286.
Butterfield, N.J. 1995. Secular distribution of Burgess-Shale-type preservation.
Lethaia, 28, 1–13.
Caron, J.B. & Jackson, D.A. 2006. Taphonomy of the Greater Phyllopod Bed
community, Burgess Shale. Palaios, 21, 451–465.
Cave, R. 1979. Sedimentary environments of the basinal Llandovery of mid-Wales.
In: Harris, A.L., Holland, C.H. & Leake, B.E. (eds) The Caledonides of
the British Isles—Reviewed. Geological Society, London, Special Publica-
tions, 8, 517–526.
Conway Morris, S. 1985. The Middle Cambrian metazoan Wiwaxia corrugata
(Matthew) from the Burgess Shale and Ogygopsis Shale, British Columbia,
Canada. Philosophical Transactions of the Royal Society of London, Series B,
307, 507–586.
Conway Morris, S. 1986. The community structure of the Middle Cambrian
Phyllopod Bed Burgess Shale. Palaeontology, 29, 423–467.
Davies, J.R., Fletcher, C.J.N., Waters, R.A., Wilson, D., Woodhall, D.G. &
Zalasiewicz, J.A. 1997. Geology of the country around Llanilar and
Rhayader. Memoir of the British Geological Survey, Sheets 178 and 170,
England and Wales.
Fletcher, T.P. & Collins, D.C. 1998. The Middle Cambrian Burgess Shale and
its relationship to the Stephen Formation in the southern Canadian Rocky
Mountains. Canadian Journal of Earth Sciences, 35, 413–436.
Received 6 February 2007; revised typescript accepted 14 June 2007.
Scientific editing by Howard Falcon-Lang
Gaines, R.R., Kennedy, M.J. & Droser, M.L. 2005. A new hypothesis for organic
preservation of Burgess Shale taxa in the Middle Cambrian Wheeler
Formation, House Range, Utah. Palaeogeography, Palaeoclimatology,
Palaeoecology, 220, 193–205.
Gostlin, K.E. & Miall, A.D. 2005. Sedimentology of the Burgess Shale: refining
the conditions for soft-bodied fossil preservation. Expanded Abstracts,
Geological Society of America, Annual Meeting, Salt Lake City, 547.
Lowe, D.R. & Guy, M. 2000. Slurry-flow deposits in the Britannia Formation
(Lower Cretaceous), North Sea: a new perspective on the turbidity current
and debris flow problem. Sedimentology, 47, 31–70.
Lowe, D.R., Guy, M. & Palfrey, A. 2003. Facies of slurry-flow deposits,
Britannia Formation (Lower Cretaceous), North Sea: implications for flow
evolution and deposit geometry. Sedimentology, 50, 45–80.
McCave, I.N. & Jones, K.P.N. 1988. Deposition of ungraded muds from high-
density non-turbulent turbidity currents. Nature, 333, 250–252.
Mulder, T. & Alexander, J. 2001. The physical character of subaqueous
sedimentary density flows and their deposits. Sedimentology, 48, 269–299.
Orr, P.J., Briggs, D.E.G. & Kearns, S.L. 1998. Cambrian Burgess Shale animals
replicated in clay minerals. Science, 281, 1173–1175.
Orr, P.J., Benton, M.J. & Briggs, D.E.G. 2003. Post-Cambrian closure of the
deep-water slope-basin taphonomic window. Geology, 31, 769–772.
Pickerill, R.K. 1994. Exceptional fossil record: Distribution of soft-tissue
preservation through the Phanerozoic: Comment and reply. Geology, 22, 183–
184.
Piper, D.J.W. 1972. Sediments of the Middle Cambrian Burgess Shale. Lethaia, 5,
169–175.
Piper, D.J.W. 1978. Turbidite muds and silts on deepsea fans and abyssal plains.
In: Stanley, D.J. & Kelling, G. (eds) Sedimentation in Submarine
Canyons, Fans and Trenches. Dowden, Hutchinson & Ross, Stroudsburg, PA,
163–176.
Powell, W.G. 2003. Greenschist-facies metamorphism of the Burgess Shale and
its implications for models of fossil formation and preservation. Canadian
Journal of Earth Sciences, 40, 13–25.
Powell, W.G., Johnston, P.A. & Collom, C.J. 2003. Geochemical evidence for
oxygenated bottom waters during deposition of fossiliferous strata of the
Burgess Shale Formation. Palaeogeography, Palaeoclimatology, Palaeoecol-
ogy, 201, 249–268.
Singer, L. & Muller, G. 1983. Diagenesis in argillaceous sediments. In: Larsen,
G. & Chilingar, G.E. (eds) Diagenesis in Sediments and Sedimentary Rocks,
2. Developments in Sedimentology, 25B, 115–212.
Stow, D.A.V. 1979. Distinguishing between fine-grained turbidites and contourites
on the Nova Scotian deep water margin. Sedimentology, 26, 371–387.
Stow, D.A.V., Faugères, J.-C., Howe, J.A., Pudsey, C.J. & Viana, A.R. 2002.
Bottom currents, contourites and deep-sea sediment drifts: current state-of-
the-art. In: Stow, D.A.V., Pudsey, C.J., Howe, J.A., Faugères, J.-C. &
Viana, A.R. (eds) Deep-water Contourite Systems; Modern Drifts and
Ancient Series, Seismic and Sedimentary Characteristics. Geological Society,
London, Memoirs, 22, 7–20.
Walcott, C.D. 1912. Cambrian of the Kicking Horse Valley, B.C.. Summary
report of the Geological Survey Branch, Department of Mines, Canada for
1911. Sessional Paper, 26, 188–191.
Whittington, H.B. 1975. The enigmatic animal Opabinia regalis, Middle
Cambrian, Burgess Shale, British Columbia. Philosophical Transactions of
the Royal Society of London, Series B, 271, 2–43.
Whittington, H.B. 1980. Exoskeleton, moult stage, appendage morphology and
habits of the Middle Cambrian trilobite Olenoides serratus. Palaeontology,
23, 171–204.
Whittington, H.B. 1985. The Burgess Shale. Yale University Press, London.
Wignall, P.B. & Newton, R. 1998. Pyrite framboid diameter as a measure of
oxygen deficiency in ancient mudrocks. American Journal of Science, 298,
537–552.
Zalasiewicz, J.A. 2001. Graptolites as constraints on models of sedimentation
across Iapetus: a review. Proceedings of the Geologists’ Association, 112,
237–251.