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 Department of Geology, University of Leicester, Leicester LE1 7RH, UK (e-mail: 2 26 Belvedere Blvd, Toronto, Ontario, M8X1K1, Canada
Abstract: We provide the most detailed sedimentological log to date through the Phyllopod Bed of the midCambrian 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 coarsergrained, 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 unit in the world. Since its discovery by Walcott in 1909 and particularly since the detailed re-descriptions of the faunas in the 1980s it has served as a key example of both the preservation of soft-bodied faunas and the diversity of life soon after the Cambrian evolutionary ‘explosion’. Non-mineralized Burgess Shale fossil remains are principally composed of kerogenized films (Butterfield 1990), sometimes with associated aluminosilicates that according to Orr et al. (1998) replicated decay-prone tissues prior to decomposition. The sedimentology of the Burgess Shale has received relatively 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 Lagerstatten are by ¨ definition rare, other factors to explain the exceptional preservation 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 preponderance 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 arthropod cuticle and other reasonably recalcitrant organic tissues (Butterfield 1995). Others (e.g. Allison & Briggs 1991, 1993; Orr et al. 2003) have suggested that after the Cambrian the increase in the amount and complexity of bioturbation eliminated the deep-water low-oxygen taphonomic window, where characteristically Burgess Shale-type preservation is found. Gaines et al. (2005) accounted for Burgess Shale-type preservation in the Middle Cambrian Wheeler Formation of Utah through a combination of influences that reduced sediment permeability, and thus

oxidant flux, to the extent that microbial decomposition was severely restrained. As well as restricted bioturbation owing to near-bottom water anoxia, low permeability was thought to be effected through deflocculated clays, early precipitation of poreoccluding carbonate cements and an absence of coarse grains such as silts, microfossils, bioclasts and faecal pellets (Gaines et al. 2005).

Fig. 1. Schematic representation of the stratigraphic relationships between the platform deposits of the Cathedral Limestone and Stephen Formations and the basinal deposits of the Burgess Shale Formation (data compiled from Fletcher & Collins 1998). The Burgess Shale Formation contains 10 members including the Walcott Quarry Shale Member, which comprises the Greater Phyllopod Bed and the interval studied herein, the Phyllopod Bed. WLM, Wash Limestone Member; GB, ‘Ginger Bed’; GML, ‘Great Marrella layer’; GEL, ‘Great Eldonia layer’.


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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.



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 righthand side of the graphic log indicates features such as pyrite lenses, shell fragments and bedding structures.

Previous sedimentological studies
The Phyllopod Bed has been described as comprising sharpbased units of calcareous siltstone grading up via interlamination into mudstone by Piper (1972), who interpreted these units to be

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


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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), overlain by alternating laminae of mudstone and well-packed calcareous siltstone, with a decrease upwards in the thickness, coarseness and packing of the siltstones, and an increase in the

thickness of mudstone laminae; upwards in each unit, mudstone laminae were described as alternating with carbon-rich laminae including some calcareous silt-sized clasts, this grading into the highest part of the unit, comprising mudstone with a small



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 massive 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 inconsistent 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-generated back-currents swept sediment into the basin.

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 between two end-members. (a) The first end-member is a fine-grained homogeneous mudstone facies (Figs 3a and 4a–d) that originally was dominantly 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 carbonatecemented 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 succession 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

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 sedimentological 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–


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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 beddingparallel even in mudstone that otherwise appears perfectly homogeneous, 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 (before significant compactional dewatering began), a broad relatively permeable zone of redox contrasts (i.e. between sediment

particles and seawater) that led first to initial ‘random’ precipitation 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 concentration 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 contourites). 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 nepheloid 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 Palaeozoic facies. Similarly, although our idealized log (Fig. 8) superficially 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.


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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-continuous 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 postcompaction).

In terms of the Bouma model the single intervals most closely compare with units Td (the millimetre-scale coarse–fine intergradations) 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.


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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 sediment 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 Bouma 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 transported 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 homogeneous 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 turbulence 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 decimetre- to metre-scale thicknesses of sediment. The entombed animals would be prevented from floating away as they accumulated 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



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 deepwater 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 timeaveraged 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 mudstone 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 preslide 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 (currentaligned) 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

(e.g. Allison & Brett 1995; Powell et al. 2003; Caron & Jackson 2006) suggests a sea floor that was at least intermittently 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 Lagerstatte 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 sedimentation 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 deposition (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 recrystallization on the fissility. As these rocks were not isoclinally folded prior to the Mesozoic Laramide Orogeny, and as overburden (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 concentration of Lagerstatten around continental margin and shelf-basin ¨ environments in the Cambrian (Allison & Briggs 1993). The perceived high relative abundance of Cambrian Lagerstatten 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 eliminated 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,


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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. Garcıa-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.

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Received 6 February 2007; revised typescript accepted 14 June 2007. Scientific editing by Howard Falcon-Lang