Robert R. Gaines

Formation of the ‘Great Unconformity’ as a trigger for the Cambrian explosion

The transition between the Proterozoic and Phanerozoic eons, beginning 542 million years (Myr) ago, is distinguished by the diversification of multicellular animals and by their acquisition of mineralized skeletons during the Cambrian period. Considerable progress has been made in documenting and more precisely correlating biotic patterns in the Neoproterozoic–Cambrian fossil record with geochemical and physical environmental perturbations, but the mechanisms responsible for those perturbations remain uncertain. Here we use new stratigraphic and geochemical data to show that early Palaeozoic marine sediments deposited approximately 540–480 Myr ago record both an expansion in the area of shallow epicontinental seas and anomalous patterns of chemical sedimentation that are indicative of increased oceanic alkalinity and enhanced chemical weathering of continental crust. These geochemical conditions were caused by a protracted period of widespread continental denudation during the Neoproterozoic followed by extensive physical reworking of soil, regolith and basement rock during the first continental-scale marine transgression of the Phanerozoic. The resultant globally occurring stratigraphic surface, which in most regions separates continental crystalline basement rock from much younger Cambrian shallow marine sedimentary deposits, is known as the Great Unconformity. Although Darwin and others have interpreted this widespread hiatus in sedimentation on the continents as a failure of the geologic record, this palaeogeomorphic surface represents a unique physical environmental boundary condition that affected seawater chemistry during a time of profound expansion of shallow marine habitats. Thus, the formation of the Great Unconformity may have been an environmental trigger for the evolution of biomineralization and the ‘Cambrian explosion’ of ecologic and taxonomic diversity following the Neoproterozoic emergence of animals.

Cambrian Burgess Shale–type deposits share a common mode of fossilization

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.

Mechanism for Burgess Shale-type preservation

Exceptionally preserved fossil biotas of the Burgess Shale and a handful of other similar Cambrian deposits provide rare but critical insights into the early diversification of animals. The extraordinary preservation of labile tissues in these geographically widespread but temporally restricted soft-bodied fossil assemblages has remained enigmatic since Walcott’s initial discovery in 1909. Here, we demonstrate the mechanism of Burgess Shale-type preservation using sedimentologic and geochemical data from the Chengjiang, Burgess Shale, and five other principal Burgess Shale-type deposits. Sulfur isotope evidence from sedimentary pyrites reveals that the exquisite fossilization of organic remains as carbonaceous compressions resulted from early inhibition of microbial activity in the sediments by means of oxidant deprivation. Low sulfate concentrations in the global ocean and low-oxygen bottom water conditions at the sites of deposition resulted in reduced oxidant availability. Subsequently, rapid entombment of fossils in fine-grained sediments and early sealing of sediments by pervasive carbonate cements at bed tops restricted oxidant flux into the sediments. A permeability barrier, provided by bed-capping cements that were emplaced at the seafloor, is a feature that is shared among Burgess Shale-type deposits, and resulted from the unusually high alkalinity of Cambrian oceans. Thus, Burgess Shale-type preservation of soft-bodied fossil assemblages worldwide was promoted by unique aspects of early Paleozoic seawater chemistry that strongly impacted sediment diagenesis, providing a fundamentally unique record of the immediate aftermath of the “Cambrian explosion.”

The Fezouata fossils of Morocco; an extraordinary record of marine life in the Early Ordovician

The discovery of the Fezouata biota in the latest Tremadocian of southeastern Morocco has significantly changed our understanding of the early Phanerozoic radiation. The shelly fossil record shows a well-recognized pattern of macroevolutionary stasis between the Cambrian Explosion and the Great Ordovician Biodiversification Event, but the rich soft-bodied Fezouata biota paints a different evolutionary picture. The Fezouata assemblage includes a considerable component of Cambrian holdovers alongside a surprising number of crown group taxa previously unknown to have evolved by the Early Ordovician. Study of the Fezouata biota is in its early stages, and future discoveries will continue to enrich our view of the dynamics of the early Phanerozoic radiation and of the nature of the fossil record. Supplementary material: A complete faunal list is available at http://www.geolsoc.org.uk/SUP18843.

A new Burgess Shale–type assemblage from the “thin” Stephen Formation of the southern Canadian Rockies

A new Burgess Shale–type assemblage, from the Stephen Formation of the southern Canadian Rocky Mountains, is described herein. It occurs near Stanley Glacier in Kootenay National Park, 40 km southeast of the type area near Field, British Columbia. While at least a dozen Burgess Shale localities are known from the “thick” Stephen Formation, the Stanley Glacier locality represents the first discovery of Burgess Shale–type fossils from the “thin” Stephen Formation. The Cathedral Escarpment, an important regional paleotopographic feature, has been considered important to the paleoecologic set- ting and the preservation of the Burgess Shale biota. However, the Stanley Glacier assemblage was preserved in a distal ramp setting in a region where no evidence of an escarpment is present. The low- diversity assemblage contains eight new soft-bodied taxa, including the anomalocaridid Stanleycaris hirpex n. gen., n. sp. (new genus, new species). Pelagic or nektobenthic predators represent the most diverse group, whereas in relative abundance, the assemblage is dominated by typical Cambrian shelly benthic taxa. The low diversity of both the benthic taxa and the ichnofauna, which includes diminutive trace fossils associated with carapaces of soft-bodied arthropods, suggests a paleoenvironment with restrictive conditions. The Stanley Glacier assemblage expands the temporal and geographic range of the Burgess Shale biota in the southern Canadian Rockies, and suggests that Burgess Shale–type assemblages may be common in the “thin” Ste- phen Formation, which is regionally widespread.

Burgess shale-type biotas were not entirely burrowed away

Burgess Shale–type biotas occur globally in the Cambrian record and offer unparalleled insight into the Cambrian explosion, the initial Phanerozoic radiation of the Metazoa. Deposits bearing except ...

Preservation of Giant Anomalocaridids in Silica-Chlorite Concretions from the Early Ordovician of Morocco

Abstract:— The recently discovered Fezouata Biota, from the Early Ordovician (late Tremadocian to late Floian) of Morocco, preserves a diverse soft-bodied fauna. While preservation is mostly of Burgess Shale-type, giant anomalocaridids also occur in siliceous concretions. Petrographic and geochemical analyses of these concretions reveal their growth history and the circumstances that led to the fossilization of nonbiomineralized anatomy within them. The large (>1 m) concretions are homogeneous in composition and geochemical characteristics, suggesting rapid, pervasive growth of mineral frameworks during decay of the large animals at, or near, the sediment-water interface. Concretions are comprised of ultrafine-grained (2–20 µm) authigenic quartz, Fe chlorite, and calcite, a composition unlike other described marine concretions. Abundant pyrite, now represented by oxide pseudomorphs, grew adjacent to the anomalocaridid carcasses, but rarely within the matrix of the concretions. This distribution indicates that sulfate reduction around the carcasses was vigorous within otherwise organic-poor sediments resulting in the establishment of prominent chemical gradients around the giant anomalocaridids that led to early precipitation of mineral overgrowth around nonbiomineralized tissues. Rapid precipitation of intergrown silica and Fe chlorite required an abundant source of silica, iron, and aluminum. These ions were likely derived from dissolution of volcanic ash in the sediments. Limited intergrown calcite (&dgr;13C avg. −12.2‰, n  =  23) precipitated from bicarbonate that was generated largely by sulfate reduction of organic tissues of the carcasses. Whereas Burgess Shale-type preservation of fossils in the Fezouata biota required suppression of degradation, exceptional preservation of anomalocaridids within the siliceous concretions resulted from extensive microbial decomposition of a large volume of organic tissues. Rapid mineralization was facilitated by localization of microbial activity around the large carcasses and must have required an unusually reactive sediment composition.

Paleoredox and pyritization of soft-bodied fossils in the ordovician frankfort shale of New York

Multiple beds in the Frankfort Shale (Upper Ordovician, New York State), including the original “Beechers Trilobite Bed,” yield fossils with pyritized soft-tissues. A bed-by-bed geochemical and sedimentological analysis was carried out to test previous models of soft-tissue pyritization by investigating environmental, depositional and diagenetic conditions in beds with and without soft-tissue preservation. Highly-reactive iron (FeHR), total iron (FeT), δ34S, organic carbon and redox-sensitive trace elements were measured. In particular, the partitioning of highly-reactive iron between iron-carbonates (Fe-carb), iron-oxides (Fe-ox), magnetite (Fe-mag), and pyrite (FeP) was examined. Overall, the multi-proxy sedimentary geochemical data suggest that the succession containing pyritized trilobite beds was deposited under a dysoxic water-column, in agreement with the paleontological data. The data do not exclude brief episodes of water-column anoxia characterized by a ferruginous rather than an euxinic state. However, the highest FeHR/FeT values and redox-sensitive trace element enrichments occur in siltstone portions of turbidite beds and in concretions, suggesting that subsequent diagenesis had a significant effect on the distribution of redox-sensitive elements in this succession. Moderately high FeHR/FeT and FeP/FeHR, low organic carbon, enriched δ34S, and the frequent presence of iron-rich carbonate concretions in beds with soft tissue preservation confirm that pyritization was favored where pore-waters were iron-dominated in sediments relatively poor in organic carbon.

Paleoecology of the olenid Trilobite Triarthrus: New evidence from Beecher's Trilobite Bed and other sites of pyritization

ABSTRACT Olenid trilobites are characteristic of low-oxygen environments in the early Paleozoic, and researchers have proposed that olenids may have harbored chemoautotrophic symbionts, allowing them to live in borderline sulfidic environments. Beds with soft-tissue preservation at the Beechers Trilobite Bed site in the Frankfort Shale and the Martin Quarry in the Whetstone Gulf Formation (both Ordovician, New York State) are dominated by the olenid Triarthrus. A bed-by-bed analysis of the sedimentology, taphonomy, paleoecology, and ichnology demonstrates that the exceptionally preserved organisms did not undergo extensive transport, and that the intervals bearing Triarthrus accumulated predominantly in the lower part of the dysaerobic zone. These intervals contain a low-diversity benthic fauna occurring in relatively low abundance, and consisting primarily of small brachiopods and trilobites. The taphonomy, in particular localized pyritization, the associated fauna, and the distribution of Triarthrus elsewhere in the Taconic foreland basin demonstrate that the environments in which Triarthrus lived were not sulfidic, and that these trilobites were unlikely to have adopted a chemoautotrophic mode of life.

Taphonomy and depositional setting of the Burgess Shale Tulip Beds, Mount Stephen, British Columbia

ABSTRACT Burgess Shale–type deposits represent exceptional preservational windows for examining the biodiversity and ecological structure of some of the earliest metazoan communities that evolved during the Cambrian Explosion. While much attention has been paid to the original Burgess Shale locality, the Walcott Quarry on Fossil Ridge, temporal and regional variations of the depositional environment of the Burgess Shale biota as a whole are still poorly understood. Here we present the first comprehensive taphonomic and sedimentological study of the Tulip Beds on Mount Stephen (Campsite Cliff Shale Member, Burgess Shale Formation), based on a time-averaged assemblage of nearly 10,000 specimens. The taphonomic characteristics—size sorting, resistance to decay, and potential flow alignment—and mode of deposition of this assemblage are compared specifically to those of the nearby and stratigraphically younger Walcott Quarry assemblage. Like other Burgess Shale–type deposits, the Tulip Beds consist of millimeter-laminated, event-derived claystone, but lack the thicker claystone layers and prominent carbonate interbeds that occur in the Walcott Quarry. These differences suggest a depositional environment lower in energy and possibly more distal to the Cathedral Escarpment. Overall, taphonomic analyses suggest no significant decay biases, transport, or sorting of the assemblage, and most specimens, benthic taxa in particular, appear to have been buried close to their living environments. Single bedding planes with large accumulations dominated by a single taxon, e.g., isolated claws of Anomalocaris, suggest short time-averaged assemblages with limited background sedimentation. Overall the Tulip Beds locality is environmentally and taphonomically comparable to the Walcott Quarry and biotic variations between the two sites are likely to be primary in nature, thus paving the way for more detailed paleoecological investigations in the future.