Burgess Shale-type Preservation Research Articles

Life in the Cambrian shallows: Exceptionally preserved arthropod and mollusk microfossils from the early Cambrian of Sweden

Abstract Burgess Shale–type (BST) Lagerstätten record an exceptional variety of Cambrian soft-bodied fauna, yet these deposits are typically restricted to outboard depositional settings >1000 km from the paleocoastline. For shallow, well-oxygenated shelf environments, our knowledge of non-mineralized animals (the majority of diversity) is severely limited, giving rise to substantial bias in our perception of Cambrian biotas. An alternate means of detecting soft-bodied Cambrian fauna, independent of paleobathymetry, is to use acid maceration to extract microscopic organic remains of non-mineralized animals, known as “small carbonaceous fossils” (SCFs). Here, a hitherto unknown diversity of Cambrian arthropod and mollusk remains are reported from shallow-marine sediments (Cambrian Stage 3 Mickwitzia Sandstone, Sweden). These microfossils comprise a variety of arthropod cuticles preserving sub-micron-scale anatomy alongside abundant radular mouthparts from mollusks—among the oldest known arthropod and molluscan SCFs on record. Significantly, at least three distinct types of fossil radula are identifiable (uniseriate, distichous, and polystichous forms), revealing that substantial diversification of the basic molluscan radula had already taken place by the early Cambrian. These cryptic elements of the biota—otherwise undetectable in such deposits—offer novel insights into Cambrian primary consumers as well as aspects of the fauna that are absent from deeper-water BST deposits.

The lower Cambrian Cranbrook Lagerstätte of British Columbia

Discovered over a century ago, the lower Cambrian (Series 2, Stage 4) Cranbrook Lagerstätte of southeastern British Columbia's Eager Formation is one of the oldest Burgess Shale-type deposits in North America. This Konservat-Lagerstätte is rich in olenelloid trilobites, but also yields a very low-diversity soft-bodied fossil assemblage including Tuzoia and Anomalocaris , and a low-diversity ichnofauna. Its scientific study, however, remains limited. A 2015 field-based investigation by the Royal Ontario Museum has revealed new information about the site's biota, depositional environment and taphonomic conditions. Not only is the Cranbrook Lagerstätte significant for early Cambrian biostratigraphy and comparisons with other Burgess Shale-type deposits, it also reveals some of the little-known diversity of life in a distal outer shelf environment during the Cambrian period. Supplementary material: Three supplementary figures (historical map, Tuzoia outlines, Anomalocaris specimens), supplementary material and methods, and a supplementary data file (generic presence/absence matrix) are available at https://doi.org/10.6084/m9.figshare.c.6850810

New soft-bodied panarthropods from diverse Spence Shale (Cambrian; Miaolingian; Wuliuan) depositional environments

AbstractThe Cambrian (Miaolingian; Wuliuan) Spence Shale Lagerstätte of northern Utah and southern Idaho is one of the most diverse Burgess Shale-type deposits of Laurentia. It yields a diverse fauna consisting of abundant biomineralized and locally abundant soft-bodied fossils, along a range of environments from shallow-water carbonates to deep-shelf dark shales. Panarthropods are the dominant component throughout the deposit, both in time and space, but whereas the trilobites and agnostoids are abundant, most of the soft-bodied taxa are only known from very few specimens. Additionally, the knowledge of soft-bodied panarthropods is currently largely limited to locations in the Wellsville Mountains of northeastern Utah. This contribution describes 21 new soft-bodied panarthropods from six locations, including the first occurrences of soft-bodied panarthropods in the High-Creek, Smithfield Creek, Spence Gulch, and Two-Mile Canyon localities. Additionally, we report the presence of bradoriids— i.e., Branchiocaris pretiosa Resser, 1929, Perspicaris? dilatus Robison and Richards, 1981, Naraoia? sp. indet., Thelxiope cf. T. palaeothalassia Simonetta and Delle Cave, 1975, and Tuzoia guntheri Robison and Richards, 1981—for the first time from the Spence Shale Lagerstätte; the first reported occurrence outside of the Burgess Shale for Thelxiope cf. T. palaeothalassia; and the first Wuliuan occurrence of Tuzoia guntheri. We also report on a new hurdiid carapace element and additional specimens of Buccaspinea cooperi? Pates et al., 2021, Dioxycaris argenta Walcott, 1886, Hurdia sp. indet., and Tuzoia retifera Walcott, 1912. This new material improves our understanding of the panarthropod fauna of the Spence Shale Lagerstätte and substantially increases our understanding of the distribution of the described taxa in time and space.

The Cambrian cirratuliform Iotuba denotes an early annelid radiation.

The principal animal lineages (phyla) diverged in the Cambrian, but most diversity at lower taxonomic ranks arose more gradually over the subsequent 500 Myr. Annelid worms seem to exemplify this pattern, based on molecular analyses and the fossil record: Cambrian Burgess Shale-type deposits host a single, early-diverging crown-group annelid alongside a morphologically and taxonomically conservative stem group; the polychaete sub-classes diverge in the Ordovician; and many orders and families are first documented in Carboniferous Lagerstätten. Fifteen new fossils of the 'phoronid' Iotuba (=Eophoronis) chengjiangensis from the early Cambrian Chengjiang Lagerstätte challenge this picture. A chaetal cephalic cage surrounds a retractile head with branchial plates, affiliating Iotuba with the derived polychaete families 'Flabelligeridae' and Acrocirridae. Unless this similarity represents profound convergent evolution, this relationship would pull back the origin of the nested crown groups of Cirratuliformia, Sedentaria and Pleistoannelida by tens of millions of years-indicating a dramatic unseen origin of modern annelid diversity in the heat of the Cambrian 'explosion'.

Aluminium-normalised trace-element paleoredox proxies and their application to the study of the conditions of Burgess Shale-type preservation

The paleo-ecological settings of Burgess Shale-type (BST) fossils have been the subject of extensive investigations, particularly with regard to the role of anoxia in their exceptional preservation. Our understanding of the complexity of both paleontological and geochemical environmental proxies has evolved with increased knowledge of niche communities and marine geochemical and biogeochemical processes. Accordingly, dated approaches should be reevaluated. The commonly applied paleoredox proxies V/Cr, Ni/Co, and V/(V + Ni), as well as the redox proxy V/Al2O3 and productivity proxy Ba/Al2O3, were all tested on the Wheeler Formation (Utah, USA) for which a fluctuating oxycline environment has been well established based on paleontological features. Subsequently, these proxies were applied to three additional BST fossil-bearing deposits: the Emigsville Member of the Kinzers Formation (Pennsylvania, USA), the Burgess Shale (British Columbia, CA) and the Emu Bay Shale (South Australia, AU). Their host shales lack perceptible V of marine origin, indicative of oxic conditions, in contrast to the Wheeler and brine-related strata from the Burgess Shale which include additional marine input of V in some beds, consistent with varying redox conditions. The black shales of the Longs Park Member (Kinzers) have elevated V/Al2O3 indicative of anoxic conditions. Molybdenum concentrations are relatively low in most samples, but minor variations tend to correlate with V/Al2O3 patterns. Ba/Al2O3 indicates high productivity with deposition of BST fossils in the Wheeler Formation, and diagenetic remobilisation of Ba owing to sediment anoxia in the Burgess Shale. Our findings show that redox proxy V/Al2O3 can be effectively applied to BST samples, whereas Ba/Al is a reliable proxy for low to moderate paleoproductivity levels and may be useful as a secondary or tangential proxy for redox conditions. Taphonomic processes leading to BST preservation occur in a variety of environments characterised by oxyclines, matgrounds, variable sedimentation rates, hypersalinity and/or anoxia. KEY POINTS V/Al2O3 and Ba/Al2O3 are effective proxies for anoxia and productivity in BST deposits. BST preservation can occur in a variety of depositional environments. Marine anoxia is not required for BST preservation.

Probability-based preservational variations within the early Cambrian Chengjiang biota (China).

The Chengjiang biota (Yunnan Province, China) is a treasure trove of soft-bodied animal fossils from the earliest stages of the Cambrian explosion. The mechanisms contributing to its unique preservation, known as the Burgess Shale-type preservation, are well understood. However, little is known about the preservation differences between various animal groups within this biota. This study compares tissue-occurrence data of 11 major animal groups in the Chengjiang biota using a probabilistic methodology. The fossil-based data from this study is compared to previous decay experiments. This shows that all groups are not equally preserved with some higher taxa more likely to preserve soft tissues than others. These differences in fossil preservation between taxa can be explained by the interaction of biological and environmental characteristics. A bias also results from differential taxonomic recognition, as some taxa are easily recognized from even poorly preserved fragments while other specimens are difficult to assign to higher taxa even with exquisite preservation.

New Cambrian vermiform organisms from Burgess Shale-type deposits of the western United States

New Cambrian vermiform organisms from Burgess Shale-type deposits of the western United States

Preservation and diagenesis of soft-bodied fossils and the occurrence of phosphate-associated rare earth elements in the Cambrian (Wuliuan) Spence Shale Lagerstätte

The Spence Shale Lagerstätte is a middle Cambrian (Miaolingian Series; Wuliuan Stage) fossil deposit in northern Utah and southern Idaho, USA. At present, it is known to preserve 89 species, from at least 10 phyla, of biomineralizing and soft-bodied taxa, and represents the only major Lagerstätte of Wuliuan age in Laurentia outside the Canadian Rockies. In addition to taxonomic diversity, the Spence Shale exposures represent a variety of depositional environments along a continental shelf, which have been affected by several diagenetic events, making it a compelling setting to improve our understanding of taphonomic and diagenetic pathways as they affect the broader Burgess Shale-type (BST) preservation. We present analyses of 28 specimens from six localities using scanning electron microscopy (SEM) and integrated energy dispersive X-ray spectrometry (EDS), together with comparisons of published data. Taxa are predominantly preserved as a combination of carbonaceous compressions, iron oxide and pyrite replacement, but eldoniids of the genus Eldonia consistently show enrichment in rare earth elements (REE). The REE, predominantly lanthanum and cerium, are enriched in the phosphate mineral monazite, which occurs in the inner coil of Eldonia ludwigi . This pattern, seen in all sampled Eldonia , differs from previous reports of REE association in BST preservation in being limited to one tissue type within a specific identifiable taxon. REE enrichment of tissues can occur during the lifetime of an individual or during fossilization, and we present both possible pathways along with an outline of taphonomic processes leading to the preservation of soft-bodied fossils from the Spence Shale. • Main preservation modes are carbonaceous compressions and iron oxides/pyrite. • Phosphatized Eldonia ludwigi have unique REE associations. • Preservation of tissue types is biased, as is found in other Paleozoic Lagerstätten.

The Chengjiang Biota inhabited a deltaic environment

The Chengjiang Biota is the earliest Phanerozoic soft-bodied fossil assemblage offering the most complete snapshot of Earth’s initial diversification, the Cambrian Explosion. Although palaeobiologic aspects of this biota are well understood, the precise sedimentary environment inhabited by this biota remains debated. Herein, we examine a non-weathered core from the Yu’anshan Formation including the interval preserving the Chengjiang Biota. Our data indicate that the succession was deposited as part of a delta influenced by storm floods (i.e., produced by upstream river floods resulting from ocean storms). Most Chengjiang animals lived in an oxygen and nutrient-rich delta front environment in which unstable salinity and high sedimentation rates were the main stressors. This unexpected finding allows for sophisticated ecological comparisons with other Burgess Shale-type deposits and emphasizes that the long-held view of Burgess Shale-type faunas as snapshots of stable distal shelf and slope communities needs to be revised based on recent sedimentologic advances.

The soft-bodied biota of the Cambrian Series 2 Parker Quarry Lagerstätte of northwestern Vermont, USA

AbstractA non-biomineralized arthropod,Protocaris marshi, was described from the lower Cambrian (Dyeran Series 2, Stage 4) of Parker's Cobble in northwestern Vermont in 1884. It represents the first fossil exhibiting Burgess Shale-type preservation to have been discovered. The locality was presumed to have been worked out and was not collected in a significant way for more than 100 years. Rediscovery of productive layers has yielded soft-bodied and lightly sclerotized taxa new to the locality, including the algaFuxianospora, a possible priapulid, a radiodont, and a specimen tentatively assigned toHerpetogaster. New specimens of the spongeLeptomitus zitteli, the bivalved arthropodTuzoia, and the chordateEmmonsaspis cambrensisprovide additional information on those taxa, and multiple specimens allow a bivalved arthropod,Vermontcaris montcalminew genus, new species, to be described. The primary mode of fossil preservation is as carbonaceous compressions. The Parker Quarry Lagerstätte complements the Kinzers Formation of Pennsylvania (also Series 2, Stage 4) in revealing the diversity of soft-bodied taxa on the southern margin of the paleocontinent Laurentia.UUID:http://zoobank.org/2eb3ae7c-6579-403e-a9ab-654c05af4a1b

Evidence for microbially mediated silver enrichment in a middle Cambrian Burgess Shale-type deposit, Mackenzie Mountains, northwestern Canada

The Selwyn basin and Mackenzie platform of northwestern Canada house an array of mineral deposits and prospects that are rich in silver, including Neoproterozoic red-bed or Kupferschiefer-type Cu and lower Paleozoic sedimentary exhalative (SEDEX) and Zn–Pb deposits. Within this overall metallogenic setting, the middle Cambrian (Drumian) Rockslide Formation was deposited under a largely oxic water column on the platform-to-basin slope along the eastern side of the Selwyn basin. The formation includes an interval termed the Ravens Throat River Lagerstätte, which is a localized Burgess Shale-type calcareous mudstone about 2 m thick that preserves soft-bodied fossils. The mudstone contains comparatively large amounts of organic carbon preserved as thin carbonaceous laminae and discontinuous seams, representing benthic microbial mats, the remains of cyanobacteria and algae that were living in the water column, fecal pellets, large coprolites, and degraded animal tissues. The upper part of the Rockslide Formation, including the fossiliferous interval, contains elevated concentrations of Ag, up to 0.47 ppm. Some of the Ag in the mudstone occurs as aggregates of elemental particles ∼10 µm in size, preferentially on the carbonaceous material comprising the coprolites. This localized enrichment suggests bioaccumulation of Ag nanoparticles or Ag+ from the water column by microorganisms on the coprolites or degrading organic matter in them. The source of the Ag may have been from penecontemporaneous SEDEX metallogeny or from broadly related subsurface fluids in the Selwyn Basin that enriched the overlying seawater.

Synchrotron imagery of phosphatized eggs in Waptia cf. W. fieldensis from the middle Cambrian (Miaolingian, Wuliuan) Spence Shale of Utah

AbstractExceptionally preserved fossil eggs and embryos provide critical information regarding paleoembryogenesis, reproductive strategies, and the early ontogeny of early arthropods, but the rarity of preservation of both eggs and egg-bearing organisms in situ limits their use in detailed evolutionary developmental (evo-devo) studies. Burgess Shale-type deposits preserve rare instances of egg-bearing arthropods as carbonaceous compressions; however, the eggs are usually poorly preserved with no compelling evidence of embryos. We describe the first record of a brooding specimen of Waptia cf. W. fieldensis from the Spence Shale, a Cambrian (Wuliuan Stage) Burgess Shale-type deposit in northeastern Utah and southeastern Idaho. This is the first record of an egg-bearing arthropod from the Spence Shale and it exhibits two distinct modes of preservation among eggs within the single clutch: carbonization and phosphatization. Unlike the egg-bearing Burgess Shale specimens, many eggs of this Utah specimen are also preserved three-dimensionally. In addition, synchrotron radiation X-ray tomographic microscopy reveals internal distributions of mineral phases, along with potential remnants of the egg membrane and attachment structures, but, as in the Burgess Shale, no explicit traces of developing embryos. The distinct modes of preservation highlight the existence of diagenetic microenvironments within some eggs, but not in others during fossilization.

Central nervous system of a 310-m.y.-old horseshoe crab: Expanding the taphonomic window for nervous system preservation

Abstract The central nervous system (CNS) presents unique insight into the behaviors and ecology of extant and extinct animal groups. However, neurological tissues are delicate and prone to rapid decay, and thus their occurrence as fossils is mostly confined to Cambrian Burgess Shale–type deposits and Cenozoic amber inclusions. We describe an exceptionally preserved CNS in the horseshoe crab Euproops danae from the late Carboniferous (Moscovian) Mazon Creek Konservat-Lagerstätte in Illinois, USA. The E. danae CNS demonstrates that the general prosomal synganglion organization has remained essentially unchanged in horseshoe crabs for >300 m.y., despite substantial morphological and ecological diversification in that time. Furthermore, it reveals that the euarthropod CNS can be preserved by molding in siderite and suggests that further examples may be present in the Mazon Creek fauna. This discovery fills a significant temporal gap in the fossil record of euarthropod CNSs and expands the taphonomic scope for preservation of detailed paleoneuroanatomical data in the Paleozoic to siderite concretion Lagerstätten of marginal marine deposits.

A juvenile-rich palaeocommunity of the lower Cambrian Chengjiang biota sheds light on palaeo-boom or palaeo-bust environments.

The fossil record, including the record of Burgess Shale-type deposits, is biased towards late ontogenetic stages. Larval stages, juvenile and subadult specimens exist but are very rare and often preserved as phosphatic fossils, resulting in biased population structures. Here, we report a new Burgess Shale-type Lagerstätte from Haiyan, China. The Haiyan palaeocommunity is extraordinary in that it is rich in fossils of early and middle ontogenetic stages of various phyla, with eggs also commonly found in the studied interval. This Lagerstätte also hosts a considerable number of new taxa-many related to later biotas of Gondwana and Laurentia. We propose that the deposit may either preserve one of the earliest nurseries in the fossil record or, alternatively, records several attempted invasions. Our study highlights the complexity of biotas and their interactions in the lower Cambrian ocean and calls for a better understanding of the mechanisms responsible for the observed spatial variation of fossil community composition in the Cambrian.

Phylogenetic response of naraoiid arthropods to early–middle Cambrian environmental change

AbstractThe Cambrian Period, primarily known for animal life diversifying, experienced global extinctions. Pulses of extinction in Cambrian Series 2 are exemplified by the disappearance of archaeocyath sponges and olenelline and redlichiid trilobites. However, the effect of such extinctions on outer shelf organisms, as typify Burgess Shale‐type (BST) deposits, remains relatively unknown. The phylogeny of naraoiid arthropods, represented in BST deposits globally, has consequently been reconstructed from either side of the Series 2 – Miaolingian extinction event to evaluate the response of offshore marine organisms to Cambrian environmental perturbation. As soft anatomy is known for only a subset of naraoiid species, exoskeletal morphology has proven important. Misszhouia and Naraoia (Naraoia) are distinguished morphometrically by posterior shield length/width and anterior shield length/posterior shield length. Morphometry has also been used to strengthen the identification of some cryptic naraoiid species and revise stratigraphic ranges. A revised naraoiid phylogeny reveals Misszhouia as a monophyletic subgenus, the former genus Pseudonaraoia nests within Naraoia and is placed in synonymy, and the systematic position and status of the Subfamily Liwiinae are sensitive to character weighting. Ten species of Naraoiidae range across the Series 2 – Miaolingian boundary, all naraoiid lineages originating during the main BST window. The persistence of outer shelf naraoiids throughout the Series 2 extinctions suggests that deeper offshore marine environments were resilient to extinction during periods of environmental stress. This study provides novel empirical support for the asylum of BST communities, which may contribute to the taxonomic longevity and widespread geographic distribution of taxa in these biotas.

Aluminosilicate haloes preserve complex life approximately 800 million years ago.

Mudstone-hosted microfossils are a major component of the Proterozoic fossil record, particularly dominating the record of early eukaryotic life. Early organisms possessed no biomineralized parts to resist decay and controls on their fossilization in mudstones are poorly understood. Consequently, the Proterozoic fossil record is compromised-we do not know whether changing temporal/spatial patterns of microfossil occurrences reflect evolution or the distribution of favourable fossilization conditions. We investigated fossilization within the approximately 1000 Ma Lakhanda Group (Russia) and the approximately 800 Ma Svanbergfjellet and Wynniatt formations (Svalbard and Arctic Canada). Vertical sections of microfossils and surrounding matrices were extracted from thin sections by focused ion beam milling. Elemental mapping and synchrotron-based infrared microspectroscopy revealed that microfossils are surrounded by haloes rich in aluminium, probably hosted in kaolinite. Kaolinite has been implicated in Cambrian Burgess Shale-type (BST) fossilization and is known to slow the growth of degraders. The Neoproterozoic mudstone microfossil record may be biased to tropical settings conducive to kaolinite formation. These deposits lack metazoan fossils even though they share fossilization conditions with younger BST deposits that are capable of preserving non-mineralizing metazoans. Thus metazoans, at least those typically preserved in BST deposits, were probably absent from sedimentary environments before approximately 800 Ma.

Re-description of the Spence Shale palaeoscolecids in light of new morphological features with comments on palaeoscolecid taxonomy and taphonomy

The middle Cambrian (Miaolingian Series; Wuliuan Stage) Spence Shale of Utah and Idaho preserves a diverse assemblage of biomineralized and soft-bodied taxa. Among the rarest specimens of this fauna are palaeoscolecid worms. Until recently, only one specimen was known from the Spence Shale, the holotype specimen of Palaeoscolex ratcliffei Robison, 1969, later included in the genus Wronascolex. This specimen is preserved as part and counterpart but missing both the posterior and anterior terminations. A new specimen, discovered by Riley Smith, preserves an everted proboscis with spines. Based on new data collected using scanning electron microscopy and energy dispersive X-ray spectrometry (SEM–EDS), and optical microscopy, the species is thought to represent a new genus, Utahscolex gen. nov., rather than a species of Palaeoscolex or Wronascolex as previously suggested. The new genus differs from the other two genera in the lack of node ornamentation of the plates, as well as the absence of microplates and platelets. Based on this case study, it is recommended that future revisions of palaeoscolecid taxonomy require knowledge of plate, platelet, and microplate ornamentation, as well as the arrangement of the plates, platelets, and microplates on the cuticle. In addition to the improved morphological information provided by the new specimen, it also advances our knowledge of the taphonomic pathways in the Spence Shale and in palaeoscolecid worms in general. The preservation of the plates of the two specimens of this species differ in elemental composition and somewhat in quality. While both the holotype and new specimen show localized magnesium and phosphorus within the plates, the holotype has a substantial iron component, whereas the new specimen instead shows elevated calcium. In addition, kerogenization, pyritization, aluminosilicification, and phosphatization can be observed throughout the specimen. The preservation varies not only between the specimens, but also within, demonstrating the high variability of preservational pathways within a Burgess Shale-type deposit, and providing insights into the circumstances that lead to soft-bodied preservation in the Spence Shale.

Proclivity of nervous system preservation in Cambrian Burgess Shale-type deposits.

Recent investigations on neurological tissues preserved in Cambrian fossils have clarified the phylogenetic affinities and head segmentation in pivotal members of stem-group Euarthropoda. However, palaeoneuroanatomical features are often incomplete or described from single exceptional specimens, raising concerns about the morphological interpretation of fossilized neurological structures and their significance for early euarthropod evolution. Here, we describe the central nervous system (CNS) of the short great-appendage euarthropod Alalcomenaeus based on material from two Cambrian Burgess Shale-type deposits of the American Great Basin, the Pioche Formation (Stage 4) and the Marjum Formation (Drumian). The specimens reveal complementary ventral and lateral views of the CNS, preserved as a dark carbonaceous compression throughout the body. The head features a dorsal brain connected to four stalked ventral eyes, and four pairs of segmental nerves. The first to seventh trunk tergites overlie a ventral nerve cord with seven ganglia, each associated with paired sets of segmental nerve bundles. Posteriorly, the nerve cord features elongate thread-like connectives. The Great Basin fossils strengthen the original description—and broader evolutionary implications—of the CNS in Alalcomenaeus from the early Cambrian (Stage 3) Chengjiang deposit of South China. The spatio-temporal recurrence of fossilized neural tissues in Cambrian Konservat-Lagerstätten across North America (Pioche, Burgess Shale, Marjum) and South China (Chengjiang, Xiaoshiba) indicates that their preservation is consistent with the mechanism of Burgess Shale-type fossilization, without the need to invoke alternative taphonomic pathways or the presence of microbial biofilms.

First occurrence of the Cambrian arthropodSidneyiaWalcott, 1911 outside of Laurentia

AbstractThe arthropodSidneyiaWalcott, 1911 is a remarkable animal of the Burgess Shale biota (Cambrian Miaolingian, Wuliuan; British Columbia, Canada), which has not been confidently reported from other Cambrian Konservat-Lagerstätten. Here we report the discovery ofSidneyiacf.inexpectansfrom the Wuliuan Mantou Formation of North China, which substantially expands the known palaeogeographical distribution of this genus. Our discovery suggests thatSidneyiahad much greater dispersal ability than hitherto thought. It also confirms the presence of exceptionally preserved fossils in the Wuliuan Mantou Formation, one of the rare Burgess Shale-type deposits of North China.

THE LIMITS OF BURGESS SHALE-TYPE PRESERVATION: ASSESSING THE EVIDENCE FOR PRESERVATION OF THE BLOOD PROTEIN HEMOCYANIN IN THE BURGESS SHALE

Cambrian deposits bearing exceptional preservation of soft-bodied fossils offer unique insights into the diversity and paleobiology of the Cambrian biota. In recent years, a growing number of studies have provided evidence that highly labile tissues, such as nervous tissues, may be more common in fossils from Burgess Shale-type deposits than previously appreciated. These discoveries have provided novel insights into the anatomy and phylogeny of Cambrian animals and have also challenged the basic understanding of the limits on the resolution of anatomical information that may be captured by Burgess Shale-type preservation. Evidence used to infer primary blood chemistry was recently reported from the iconic Burgess Shale fossil Marrella splendens, the first such report from a Cambrian Lagerstatte. Diminutive crystals of a copper sulfide mineral found in association with Marrella were interpreted as evidence that the animal's blood contained the Cu-bearing protein hemocyanin. Copper minerals in shales in general, and in the Burgess Shale in particular, however, may have a complex geologic history. Using microscopy and SEM-EDS elemental mapping of material from the Walcott Quarry as well as from the Burgess Shale fossil quarry at Marble Canyon, we assess the timing of emplacement of Cu-sulfides in fossil-bearing mudrocks of the Burgess Shale. These data demonstrate that copper minerals are most conspicuously concentrated within veins that are oriented sub-perpendicular to bedding. These late-stage fluid flow paths allowed Cu-mineralizing fluids to infiltrate bedding plane-parallel cracks, which commonly run through Burgess Shale fossils and extend into the matrix around them. Based upon the relationships of mineral phases associated with fossils, it appears that Cu-mineralization occurred after metamorphic volatilization of a large proportion of the carbonaceous material that originally comprised the fossils. We find that copper minerals in the Burgess Shale are of demonstrably post-Cambrian origin, a secondary overprint that cannot be used to infer original blood chemistry or tissue composition of its fossils.