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Robert R. Gaines1, Derek E.G. Briggs 2, Zhao Yuanlong3
Geology Department, Pomona College, Claremont, California 91711, USA 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 fundamental 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 mineral 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 mechanism responsible for the preservation of BST assemblages. Early authigenic mineral replacement preserves certain anatomical features 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 geologic 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 Cambrian 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). 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 precipitation 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 Formation, 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 analyses supported a carbonaceous (Whittington, 1971) or alumino-silicate composition (Conway Morris, 1977, 1986). Organic remains of features 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 conservation 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 mudstone 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 decomposing 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 fossils 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 tissues 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 mechanism 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).
© 2008 The Geological Society of America. For permission to copy, contact Copyright Permissions, GSA, or firstname.lastname@example.org. GEOLOGY, October 2008 36; no. 10; p. 755–758; doi: 10.1130/G24961A.1; 2 ﬁgures. Geology, October 2008; v.
MATERIALS AND METHODS In this study we analyzed 53 fossils from 11 BST deposits (Appendix 1). All deposits analyzed have undergone less metamorphism than the Burgess Shale, as evidenced by a predominance of illite over chlorite 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). Element distribution was determined by production of X-ray maps at micronscale 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 preservation 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 distinguished from the matrix using EDX analysis or SEM imaging, even though they are clearly visible to the naked eye as dark areas. These specimens 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 morphology. 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; Butterﬁeld 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 compositional difference between fossil and matrix was detected, even where carbon ﬁlms are so severely degraded that they are below the threshold of
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 represents 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.
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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 augmented by authigenic mineral replication of particular features. Carbonaceous remains have also been documented in isolated elements of BST fossils extracted by HF digestion from the Mount Cap Formation (Butterﬁeld, 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 exoskeleton, 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 graptolites were shown to be replaced progressively by alumino-silicate mineral ﬁlms with increasing metamorphic grade as a result of late diagenetic processes (Page et al., 2007). These data suggest that the original composition 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 (Butterﬁeld, 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
taphonomic pathway in Chengjiang fossils suggests an enrichment of reactive iron in the associated sediment (Briggs et al., 1996). Such enrichment would be expected to result from deposition under an anoxic water column (Poulton and Canﬁeld, 2005) or near a source of terrigenous clastics, 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 concentrated 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 phosphate 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 Formation 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 authigenic 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 preservation considered here, representing different primary modes of fossilization, 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 otherwise 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 carbonaceous 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. Protection of carcasses has been attributed to clay-organic interactions (Butterﬁeld, 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 microbial decomposition (Gaines et al., 2005). Further study of the sedimentology and geochemistry of these deposits is required to determine which of these possibilities is most likely.
APPENDIX: FOSSILS ANALYZED
Figure 2. Secondary emission scanning electron microscope (SE-SEM) micrographs of carbonaceous residues comprising Burgess Shale–type (BST) fossils. A: Margin of indeterminate vermiform metazoan deﬁned by black (conductive), amorphous carbonaceous 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 represents 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.
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 indeterminate 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 Formation [MC: 1 Leanchoilia? sp. cf. protogonia (arthropod), 1 Yuknessia
GEOLOGY, October 2008
simplex (alga)]; the Spence Shale Member, Langston Formation [MC: 1 Canadaspis? (arthropod), 1 eldoniid (metazoan), 2 Wiwaxia (lophotrochozoan), 1 indeterminate metazoan cuticle, 2 indeterminate metazoans, 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 indeterminate metazoan). (3) Two Proterozoic deposits yield compression fossils: the Neoproterozoic Doushantuo Formation (Miahoe Biota) (3 indeterminate 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 electron 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 acquisition 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 problematic 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 Paleontology, v. 82, p. 238–254, doi: 10.1666/06–086.1. Budd, G.E., 1998, Arthropod body-plan evolution in the Cambrian with an example 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 threedimensional 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 explosion: 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 Palaeontology, v. 20, 155 p.
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