Cambrian Burgess Shale–type deposits share a common mode
of fossilization
Robert R. Gaines1, Derek E.G. Briggs 2, Zhao Yuanlong3
1
Geology Department, Pomona College, Claremont, California 91711, USA
2
Department of Geology and Geophysics, Yale University, New Haven, Connecticut 06520, USA
3
Key Laboratory for Paleobiology, Guizhou University, Guiyang, China
ABSTRACT
Although Cambrian Burgess Shale–type (BST) biotas are fun-
damental to understanding the radiation of metazoans, the nature
of their extraordinary preservation remains controversial. There
remains disagreement about the importance of the role of early min-
eral replication of soft tissues versus the conservation of primary
organic remains. Most prior work focused on soft-bodied fossils
from the two most important BST biotas, those of the Burgess Shale
(Canada) and Maotianshan Shale (Chengjiang, China). Fossils from
these two deposits do not provide ideal candidates for specimen-level
taphonomic study because they have been altered: the Burgess Shale
by greenschist facies metamorphism and the Maotianshan Shale by
intensive subsurface weathering. Elemental mapping of soft-bodied
fossils from 11 other BST deposits worldwide demonstrates that BST
preservation represents a single major taphonomic pathway that may
share a common cause wherever it occurs. The conservation of organic
tissues, and not early authigenic mineralization, is the primary mech-
anism responsible for the preservation of BST assemblages. Early
authigenic mineral replacement preserves certain anatomical fea-
tures of some specimens, but the preservation of non-biomineralized
BST fossils requires suppression of the processes that normally lead
to the degradation of organic remains in marine environments.
Keywords: taphonomy, Chengjiang, organic preservation, authigenic
mineralization.
INTRODUCTION
Burgess Shale–type (BST) biotas are of critical importance to
understanding the early evolution of the Metazoa (Conway Morris,
1989a; Butterfield, 2003; Briggs and Fortey, 2005). The great majority of
species preserved in BST deposits lack biomineralized tissues (Conway
Morris, 1986). Preservation of soft-bodied organisms is rare in the geo-
logic record (Briggs, 2003) and BST deposits provide a record of early
Phanerozoic biodiversity otherwise unknown. The mode of preservation
of these exceptional biotas, however, has been much debated and the
precise circumstances that facilitated exceptional preservation in BST
deposits remain disputed.
It has long been recognized that an anomalously large number
of deposits in Lower and Middle Cambrian strata yield exceptionally
preserved biotas compared to the rest of the Phanerozoic (Allison and
Briggs, 1993). Soft-bodied fossils occur in abundance at nine Cambrian
localities worldwide (Conway Morris, 1998), and more rarely in perhaps
as many as 40 other deposits of Cambrian age (e.g., Conway Morris,
1989b; Steiner et al., 2005). Although a small number of Cambrian
deposits preserve three-dimensional soft-bodied fossils by replacement
in calcium phosphate (e.g., Waloszek, 2003), the great majority of Cam-
brian exceptional biotas are preserved as two-dimensional compression
fossils. Deposits yielding biotas preserved in this characteristic manner
are termed Burgess Shale–type deposits after Walcott’s classic locality
in the Canadian Rockies (Conway Morris, 1989a, 1989b). It has been
suggested that this type of preservation is largely absent from the fossil
record after the Middle Cambrian (Butterfield, 1995).
© 2008 The Geological Society of America. For permission to copy, contact Copyright Permissions, GSA, or editing@geosociety.org.
GEOLOGY,
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2008; v. 36; no. 10; p. 755–758; doi: 10.1130/G24961A.1; 2 figures.
The importance of organic preservation versus early authigenic
mineral replacement of soft tissues in BST deposits has been disputed.
A number of microbial decomposition reactions facilitate the precipita-
tion of minerals on decaying tissues, replicating their form (Briggs, 2003).
Once soft tissues are replicated by mineral templates, they may survive to
enter the fossil record even if the original organic material is lost. On the
other hand, the preservation of whole biotas as organic remains requires
inhibition of the normal processes of decay.
Most prior work focused on the preservation of two of the most
important BST assemblages, those of the Middle Cambrian Burgess
Shale and the Lower Cambrian Maotianshan Shale (Yuanshan For-
mation, Chengjiang, China). Typically, BST fossils are preserved as
dark-colored films, often reflective, exposed on bedding surfaces of
the host mudstones. Burgess Shale fossils were first interpreted to be
the result of siliceous replacement (Walcott, 1919). Subsequent analy-
ses supported a carbonaceous (Whittington, 1971) or alumino-silicate
composition (Conway Morris, 1977, 1986). Organic remains of fea-
tures of some Burgess Shale fossils were documented in thin section
and isolated by digestion in HF (Butterfield, 1990). These findings led
to the definition of Burgess Shale–type preservation as the conserva-
tion of two-dimensional carbonaceous compressions in marine shales
(Butterfield, 1995). However, elemental mapping of Burgess Shale
fossils subsequently demonstrated that the abundance of Al, Si, and
K varies in different morphological features and relative to the mud-
stone matrix (Orr et al., 1998). This discovery led to the conclusion
that more labile tissues of Burgess Shale fossils are replicated in clay
minerals, which precipitated onto the organic templates of decom-
posing organisms in the early burial environment (Orr et al., 1998).
Orr et al. (1998) concluded that while organic remains of the more
decay-resistant cuticles are preserved, authigenic mineralization was
the principal pathway by which the more labile tissues in fossils of the
Burgess Shale were preserved.
Elemental mapping of fossils from the Maotianshan Shale
revealed that two modes of preservation are important in that deposit.
In most cases the major morphological features of Chengjiang fos-
sils are preserved as carbonaceous compressions; however, features
of many of these fossils are preserved in pyrite (Gabbott et al., 2004;
Hu, 2005). Pyrite mineralization is interpreted to have occurred soon
after burial during limited bacterial sulfate reduction around the most
labile tissues, defining their morphology; other, more recalcitrant tis-
sues are preserved as organic compressions (Gabbott et al., 2004).
Thus, analyses of fossils from the Maotianshan Shale (Gabbott et al.,
2004) and from the Burgess Shale (Orr et al., 1998) suggested that
different diagenetic pathways led to the preservation of fossils in the
two deposits. Furthermore, these results implied that no single mecha-
nism was responsible for the preservation of BST biotas. Although the
Burgess Shale and Maotianshan Shale have yielded the most important
Cambrian biotas, their fossils may not provide the most appropriate
basis for understanding BST preservation. Burgess Shale fossils from
Fossil Ridge have been altered by greenschist facies metamorphism
(Powell, 2003) and those of the Maotianshan Shale have been altered
by extensive weathering (Zhu et al., 2001).
755
MATERIALS AND METHODS
In this study we analyzed 53 fossils from 11 BST deposits (Appen-
dix 1). All deposits analyzed have undergone less metamorphism than
the Burgess Shale, as evidenced by a predominance of illite over chlo-
rite and muscovite in the matrix of mudstones, and less weathering than
Maotianshan Shale, as evidenced by higher specific gravity. A variety of
taxa without mineralized skeletons was analyzed, including arthropods,
priapulids and other worms, eldoniids, problematic taxa, and algae or
alga-like fossils. The least weathered fossils available from each deposit
were selected for analysis. Fossils were analyzed uncoated using field
emission scanning electron microscope–energy dispersive X-ray analysis
(SEM-EDX) at low voltage (5 kV) in order to minimize beam penetration.
A subset of samples was also analyzed at higher voltage (15–20 kV). Ele-
ment distribution was determined by production of X-ray maps at micron-
scale resolution, augmented by spot and line scan transects.
RESULTS
Elemental mapping revealed that all 53 fossils analyzed are preserved
as carbonaceous films or their degraded remains. Authigenic minerals are
responsible for preservation of a morphological feature (the gut) in only
one of the samples analyzed. The fossils display carbonaceous preserva-
tion that ranges from robust continuous films to what are interpreted as
degraded remains of films that are below the threshold of EDX detection.
Elemental mapping of robust and continuous films (Fig. 1) reveals
sharp contrasts in composition between the fossils and the matrix. The
carbonaceous remains are clearly defined by strong enrichment in C and
depletion of Al, Si, and O. Where present, Ca and Mg are also depleted
relative to the matrix. Under SEM magnification, these films are readily
distinguished from the crystalline claystone matrix by enhanced conduc-
tivity and amorphous habit (Fig. 2). Films vary in thickness from several
microns to <1 μm, but do not preserve original microstructures.
At the opposite end of the spectrum are fossils that cannot be dis-
tinguished from the matrix using EDX analysis or SEM imaging, even
though they are clearly visible to the naked eye as dark areas. These speci-
mens exhibit no compositional difference from the matrix and no textural
indication that authigenic minerals were once present, such as crystal
pseudomorphs or molds. Such fossils are preserved as degraded carbon
films, which define morphological features but retain insufficient carbon to
be detected; of the elements analyzed, carbon is the most difficult to detect
because of its low atomic number and tendency to emit low-energy X-ray
radiation. This conclusion is supported by analyses of individual fossils
that revealed a gradient in the nature of carbonaceous material, from
robust through weakly detectable to nondetectable, and is consistent with
previous work (Gabbott et al., 2004).
Authigenic pyrite is present in two of the fossils analyzed, as euhedral
crystals (~5 μm) and as framboids (~10 μm). In both cases, pyrite occurs
in patches at the sediment-fossil interface that do not replicate fossil mor-
phology. However, replacement by early authigenic calcium phosphate
preserves the gut of the arthropod Dicranocaris guntherorum (Briggs
et al., 2008) in three dimensions within a two-dimensional carbonaceous
film that defines the rest of the morphology of the specimen.
Authigenic mineral phases have been shown to coat organic remains
of soft-bodied fossils from the Burgess Shale (Orr et al., 1998; Butter-
field et al., 2007). However, there is no evidence that alumino-silicates or
other authigenic mineral phases are present on the surface of the fossils
analyzed here but are obscured by robust carbonaceous remains. No com-
positional difference between fossil and matrix was detected, even where
carbon films are so severely degraded that they are below the threshold of

C O Al Si Fe
A′

A
B′

B
C′

C
D′

D
Figure 1. Examples of scanning electron microscope–energy dispersive X-ray (SEM-EDX) elemental maps illustrating preservation of
Burgess Shale–type (BST) fossils as carbonaceous films (A–D) and photographs showing location on each specimen at which map data
were taken (A′–D′). Brightness of color corresponds to abundance of each element. A: Carapace of Tuzoia, Kaili Formation; scale repre-
sents 300 μm. A′: Scale represents 5 mm. B: Margin of indeterminate vermiform metazoan (left), Spence Shale (A); scale represents 400 μm.
B′: Scale represents 5 mm. C: Indeterminate alga (top), Doushantuo Formation; scale represents 100 μm. C′: Scale represents 2 mm. D: Gut
trace of indeterminate metazoan, Pioche Formation; scale represents 400 μm. D′: Scale represents 1 mm.

756 GEOLOGY, October 2008

detection. Analysis of a subset of carbonaceous samples at higher voltages
(15 kV and 20 kV) for deeper beam penetration also revealed no evidence
for mineral coatings beneath a surficial carbonaceous layer.
DISCUSSION
These results demonstrate that fossils analyzed from the 11 deposits
are preserved as carbonaceous compressions that are sometimes aug-
mented by authigenic mineral replication of particular features. Carbo-
naceous remains have also been documented in isolated elements of BST
fossils extracted by HF digestion from the Mount Cap Formation (Butter-
field, 1994), in the gut of the trilobite Olenoides from the Kaili biota (Lin,
2007), and in soft-bodied fossils from the Middle Cambrian portion of the
Pioche Formation (Moore and Lieberman, 2005).
Recently published data indicate that conservation of carbonaceous
remains was the primary process involved in preserving the fossils of the
Burgess Shale. A petrologic and microanalytical study of fossils from
the Walcott Quarry interpreted the replication of tissues in alumino-silicate
minerals as primarily resulting from late-stage metamorphic replacement
or overgrowth of carbonaceous material (Butterfield et al., 2007). These
findings, which are based on mineral phase relationships in trilobite exo-
skeleton, in mineralized arthropod guts, and in veinlets that cut soft-bodied
fossils, are supported by the demonstration of a similar phenomenon in
graptolites across a metamorphic gradient. Originally carbonaceous grap-
tolites were shown to be replaced progressively by alumino-silicate min-
eral films with increasing metamorphic grade as a result of late diagenetic
processes (Page et al., 2007). These data suggest that the original com-
position of Burgess Shale fossils was carbonaceous, consistent with the
composition of fossils from other deposits analyzed in this study.
Although pyrite has been reported in association with soft-bodied
fossils from the Burgess Shale and the Mount Cap Formation (Butter-
field, 2003; Garcia-Bellido and Collins, 2006), extensive preservation
of the outline of soft-tissues in Cambrian faunas in pyrite has only been
reported in the Maotianshan Shale (Gabbott et al., 2004). This additional
A B
C D
Figure 2. Secondary emission scanning electron microscope
(SE-SEM) micrographs of carbonaceous residues comprising
Burgess Shale–type (BST) fossils. A: Margin of indeterminate vermi-
form metazoan defined by black (conductive), amorphous carbo-
naceous film (top and left) delineated sharply from crystalline clay
matrix (lower right), Spence Shale; scale represents 20 μm. B: Beltina
(alga?) showing patches of black amorphous carbonaceous material
around margins and at center, with weakly detectable carbonaceous
remains composing interior of the fossil, Greyson Shale; scale rep-
resents 500 μm. C: Yuknessia simplex (alga), Wheeler Formation,
showing discrete flakes of carbonaceous material that compose the
fossil. D: Detail of Selkirkia (priapulid) showing thin, discontinuous
carbonaceous film that appears dark compared to bright crystalline
matrix, Wheeler Formation; scale represents 5 μm.
GEOLOGY, October 2008
taphonomic pathway in Chengjiang fossils suggests an enrichment of
reactive iron in the associated sediment (Briggs et al., 1996). Such enrich-
ment would be expected to result from deposition under an anoxic water
column (Poulton and Canfield, 2005) or near a source of terrigenous clas-
tics, or from storm-influenced remobilization of sediments in which pyrite
had formed previously (Raiswell et al., 2008). The Maotianshan Shale
is significantly enriched in reactive iron relative to other BST deposits
(Hammarlund, 2007); although it is possible that reactive iron was con-
centrated in the Maotianshan Shale as a result of recent weathering, iron
enrichment may represent the only taphonomically significant difference
between the Chengjiang and other BST deposits.
Replication of selected soft tissues in authigenic calcium phos-
phate is another important auxiliary pathway in BST deposits. Although
uncommon, three-dimensional preservation of arthropod guts in calcium
phosphate occurs in the Burgess Shale and Maotianshan Shale (Briggs,
1981; Butterfield, 2002), in addition to the example from the Wheeler For-
mation documented here. Phosphatization of the midgut glands, which
sometimes preserves subcellular details, has been linked to the abundance
of unordered calcium phosphate present in the living organ (Butterfield,
2002). Extensive three-dimensional replication of muscle-tissues in authi-
genic minerals occurs in the fossils from Sirius Passet (in silica; Budd,
1998) and the Emu Bay Shale (in calcium phosphate: Briggs and Nedin,
1997); these biotas differ taphonomically from the BST mode of preserva-
tion considered here, representing different primary modes of fossiliza-
tion, and must be considered separately from other BST deposits.
CONCLUSIONS
The analyses presented here, together with previously published data,
indicate that BST deposits share a common taphonomic pathway that led to
the preservation of fossils as carbonaceous films (Butterfield, 1995). These
results also confirm that compression fossils of nonmineralized algae in
Proterozoic shales followed the same pathway (Butterfield, 1995). Auxiliary
mineral replacements that occur in association with the carbonaceous films
have the potential to preserve more labile structures, which would other-
wise be lost (Orr et al., 1998; Butterfield, 2002). These associated mineral
precipitates may reflect differences in porewater chemistry and in sediment
composition and provide insight into the diagenetic processes under which
fossilization occurred. However, in the great majority of cases it is a carbo-
naceous film alone that defines the overall morphology of the fossils.
These results indicate that BST preservation represents a global
phenomenon that required suppression of normal processes of decay in
marine sediments. Two types of models have been proposed to explain the
preservation of BST fossils as organic remains. The first requires physical
and/or chemical protection of carcasses entombed in sediments. Protec-
tion of carcasses has been attributed to clay-organic interactions (Butter-
field, 1995), or adsorption of Fe3+ onto biopolymers (Petrovich, 2001),
both preventing the activity of enzymes involved in decomposition. The
second requires an early cessation of normal diagenetic processes, via
rapid occlusion of sediment porosity soon after deposition, resulting in
reduced flux of oxidants into the sediments and the preclusion of micro-
bial decomposition (Gaines et al., 2005). Further study of the sedimentol-
ogy and geochemistry of these deposits is required to determine which of
these possibilities is most likely.
APPENDIX: FOSSILS ANALYZED
The 53 fossils analyzed came from 11 Burgess Shale–type (BST)
deposits. (1) Five important deposits are from the Lower (LC) and Middle
Cambrian (MC): the Kaili Formation [MC: 2 Tuzoia (arthropod), 1 inde-
terminate arthropod cuticle, 3 Pararotadiscus guizhouensis (eldoniid),
1 indeterminate alga]; the Kinzers biota of the Ledger Formation
(LC: 1 indeterminate cuticle, 1 indeterminate alga); the Marjum Forma-
tion [MC: 1 Leanchoilia? sp. cf. protogonia (arthropod), 1 Yuknessia
757
simplex (alga)]; the Spence Shale Member, Langston Formation [MC:
1 Canadaspis? (arthropod), 1 eldoniid (metazoan), 2 Wiwaxia (lopho-
trochozoan), 1 indeterminate metazoan cuticle, 2 indeterminate meta-
zoans, 3 Marpolia spissa (alga), 1 Margaretia (alga)]; and the Wheeler
Formation [MC: 1 Canadaspis?, 1 Dicranocaris guntherorum (arthropod),
1 Elrathia kingii (arthropod: soft parts analyzed), 1 Selkirkia (priapulid),
2 Margaretia, 1 Marpolia spissa, 3 Morania (alga?), 2 Yuknessia simplex
(alga), 1 indeterminate vermiform metazoan or gut trace]. (2) Four
“minor” deposits are from the Lower and Middle Cambrian, and have
yielded few soft-bodied fossils: the Balang Formation (LC: 4 Morania), the
Latham Shale [LC: 1 Anomalocaris appendage (arthropod)], the Metaline
Formation (MC: 1 Margaretia), and the Pioche Formation (MC: 1 indeter-
minate metazoan). (3) Two Proterozoic deposits yield compression fossils:
the Neoproterozoic Doushantuo Formation (Miahoe Biota) (3 indetermi-
nate algae); and the Mesoproterozoic Greyson Shale [5 Beltina (alga),
3 Grypania (alga)].
ACKNOWLEDGMENTS
We thank N. Butterfield, J.-B. Caron, P. Orr, and A. Page for comments and
discussion, and D. Haley, D. Tanenbaum, C. Taylor, and Z. Jiang for scanning elec-
tron microscope assistance. Some analyses were performed by R. Goossen and
B. Markle. We thank S. Halgedahl, P. Jameson, R. Jarrard, M. Kooser, J. Scabelund,
and G. Schofield for access to collections for analysis. This work was supported by
the National Science Foundation for collaborative research between Gaines and
Briggs (grants EAR-0518732 and EAR-0518547) and for major equipment acqui-
sition by Gaines (DMR-0618417), by a D.L. and S.H. Hirsch Research Initiation
grant to Gaines, and by Chinese Natural Science Foundation grant 40672018 to
Zhao Yuanlong.
REFERENCES CITED
Allison, P.A., and Briggs, D.E.G., 1993, Exceptional fossil record: Distribution
of soft-tissue preservation through the Phanerozoic: Geology, v. 21, p. 527–
530, doi: 10.1130/0091–7613(1993)021<0527:EFRDOS>2.3.CO;2.
Briggs, D.E.G., 1981, The arthropod Odaraia alata Walcott, Middle Cambrian,
Burgess Shale, British Columbia: Royal Society of London Philosophical
Transactions, ser. B, v. 291, p. 541–582, doi: 10.1098/rstb.1981.0007.
Briggs, D.E.G., 2003, The role of decay and mineralization in the preservation of
soft-bodied fossils: Annual Review of Earth and Planetary Sciences, v. 31,
p. 275–301, doi: 10.1146/annurev.earth.31.100901.144746.
Briggs, D.E.G., and Fortey, R.A., 2005, Wonderful strife: Systematics, stem groups,
and the phylogenetic signal of the Cambrian radiation: Paleobiology, v. 31,
p. 94–112, doi: 10.1666/0094–8373(2005)031[0094:WSSSGA]2.0.CO;2.
Briggs, D.E.G., and Nedin, C., 1997, The taphonomy and affinities of the prob-
lematic fossil Myoscolex from the Lower Cambrian Emu Bay Shale of
South Australia: Journal of Paleontology, v. 71, p. 22–32.
Briggs, D.E.G., Raiswell, R., Bottrell, S.H., Hatfield, D., and Bartels, C., 1996,
Controls on the pyritization of exceptionally preserved fossils: An analysis
of the Lower Devonian Hunsrück Slate of Germany: American Journal of
Science, v. 296, p. 633–663.
Briggs, D.E.G., Lieberman, B.S., Hendricks, J.R., Halgedahl, S.L., and Jarrard,
R.D., 2008, Middle Cambrian arthropods from Utah: Journal of Paleontol-
ogy, v. 82, p. 238–254, doi: 10.1666/06–086.1.
Budd, G.E., 1998, Arthropod body-plan evolution in the Cambrian with an exam-
ple from anomalocaridid muscle: Lethaia, v. 31, p. 197–210.
Butterfield, N.J., 1990, Organic preservation of non-mineralizing organisms and
the taphonomy of the Burgess Shale: Paleobiology, v. 16, p. 272–286.
Butterfield, N.J., 1994, Burgess Shale-type fossils from a Lower Cambrian
shallow-shelf sequence in northwestern Canada: Nature, v. 369, p. 477–
479, doi: 10.1038/369477a0.
Butterfield, N.J., 1995, Secular distribution of Burgess Shale-type preservation:
Lethaia, v. 28, p. 1–13, doi: 10.1111/j.1502–3931.1995.tb01587.x.
Butterfield, N.J., 2002, Leanchoilia guts and the interpretation of three-
dimensional structures in Burgess Shale-type fossils: Paleobiology, v. 28,
p. 155–171, doi: 10.1666/0094–8373(2002)028<0155:LGATIO>2.0.CO;2.
Butterfield, N.J., 2003, Exceptional fossil preservation and the Cambrian explo-
sion: Integrative and Comparative Biology, v. 43, p. 166–177, doi: 10.1093/
icb/43.1.166.
Butterfield, N.J., Balthasar, U., and Wilson, L.A., 2007, Fossil diagenesis in
the Burgess Shale: Paleontology, v. 50, p. 537–543, doi: 10.1111/j.1475–
4983.2007.00656.x.
Conway Morris, S., 1977, Fossil priapulid worms: Special Papers in Palaeontol-
ogy, v. 20, 155 p.
758
Conway Morris, S., 1986, The community structure of the Middle Cambrian
phyllopod bed (Burgess Shale): Palaeontology, v. 29, p. 423–467.
Conway Morris, S., 1989a, Burgess Shale faunas and the Cambrian explosion:
Science, v. 246, p. 339–346, doi: 10.1126/science.246.4928.339.
Conway Morris, S., 1989b, The persistence of Burgess Shale-type faunas: Impli-
cations for the evolution of deep-water faunas: Royal Society of Edinburgh
Transactions, v. 80, p. 271–283.
Conway Morris, S., 1998, The crucible of creation: Oxford, Oxford University
Press, 242 p.
Gabbott, S.E., Hou, X.G., Norry, M.J., and Siveter, D.J., 2004, Preservation of Early
Cambrian animals of the Chengjiang biota: Geology, v. 32, p. 901–904.
Gaines, R.R., Kennedy, M.J., and 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, v. 220, p. 193–205, doi: 10.1016/j.palaeo.2004.07.034.
Garcia-Bellido, D.C., and Collins, D.H., 2006, A new study of Marrella splen-
dens (Arthropoda, Marrellomorpha) from the Middle Cambrian Burgess
Shale, British Columbia, Canada: Canadian Journal of Earth Sciences,
v. 43, p. 721–742, doi: 10.1139/E06–012.
Hammarlund, E., 2007, The ocean chemistry at Cambrian deposits with excep-
tional preservation and the influence of sulfate on soft-tissue decay [M.S.
thesis]: Odense, Denmark, University of Southern Denmark, 64 p.
Hu, S., 2005, Taphonomy and palaeoecology of the Early Cambrian Chengjiang
Biota from eastern Yunnan, China [Ph.D. thesis]: Berliner Paläobiologische
Abhandlungen 7, 189 p.
Lin, J.P., 2007, Preservation of the gastrointestinal system in Olenoides (Trilo-
bita) from the Kaili Biota (Cambrian) of Guizhou, China, in Laurie, J.R.,
ed., South Australia 2006: Papers from the XI International Conference of
the Cambrian Stage Subdivision Working Group, South Australia 2006:
Association of Australasian Palaeontologists Memoir 33, p. 179–189.
Moore, R.A., and Lieberman, B.S., 2005, Taphonomy of Lower and Middle Cam-
brian arthropods from the Pioche Shale of Nevada: Geological Society of
America Abstracts with Programs, v. 37, no. 7, p. 548.
Orr, P.J., Briggs, D.E.G., and Kearns, S.L., 1998, Cambrian Burgess Shale
animals replicated in clay minerals: Science, v. 281, p. 1173–1175, doi:
10.1126/science.281.5380.1173.
Page, A., Gabbott, S.E., and Wilby, P.R., 2007, A reassessment of fossil preserva-
tion in the Burgess Shale: Palaeontological Association Annual Meeting,
51st, Abstracts, p. 43.
Petrovich, R., 2001, Mechanisms of fossilization of the soft-bodied and lightly
armored faunas of the Burgess Shale and of some other classical localities:
American Journal of Science, v. 301, p. 683–726, doi: 10.2475/ajs.301.8.683.
Poulton, S.W., and Canfield, D.E., 2005, Development of a sequential extrac-
tion procedure for iron: Implications for iron partitioning in continentally
derived particulates: Chemical Geology, v. 214, p. 209–221, doi: 10.1016/
j.chemgeo.2004.09.003.
Powell, W., 2003, Greenschist-facies metamorphism of the Burgess Shale and
its implications for models of fossil formation and preservation: Canadian
Journal of Earth Sciences, v. 40, p. 13–25, doi: 10.1139/e02–103.
Raiswell, R., Newton, R., Bottrell, S.H., Coburn, P.M., Briggs, D.E.G., Bond,
D., and Poulton, S.W., 2008, Turbidite depositional influences on the dia-
genesis of Beecher’s Trilobite Bed and the Hunsrück Slate; sites of soft
tissue pyritization: American Journal of Science, v. 308, p. 105–129, doi:
10.2475/02.2008.01.
Steiner, M., Zhu, M., Zhao, Y., and Erdtmann, B.D., 2005, Burgess Shale-type
fossil associations of South China: Palaeoclimatology, Palaeogeography,
Palaeoecology, v. 220, p. 129–152, doi: 10.1016/j.palaeo.2003.06.001.
Walcott, C.D., 1919, Middle Cambrian algae: Smithsonian Miscellaneous Col-
lections, v. 67, p. 217–260.
Waloszek, D., 2003, The ‘Orsten’ window—A three-dimensionally preserved
Upper Cambrian meiofauna and its contribution to our understanding of
the evolution of Arthropoda: Paleontological Research, v. 7, p. 71–88, doi:
10.2517/prpsj.7.71.
Whittington, H.B., 1971, The Burgess Shale: History of research and preservation
of fossils: North American Paleontological Convention Proceedings, v. 2,
part I, p. 1170–1201.
Zhu, M., Zhang, J.M., and Li, G.X., 2001, Sedimentary environments of the Early
Cambrian Chengjiang Biota: Sedimentology of the Yu’anshan Formation
in Chengjiang County, eastern Yunnan: Acta Palaeontologica Sinica, v. 40,
p. 80–105.
Manuscript received 19 March 2008
Revised manuscript received 6 June 2008
Manuscript accepted 10 June 2008
Printed in USA
GEOLOGY, October 2008