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The middle (Wuliuan Stage) Cambrian Burgess Shale is famous for its exceptional preservation of diverse and abundant soft-bodied animals through the “thick” Stephen Formation. However, with the exception of the Walcott Quarry (Fossil Ridge) and the stratigraphically older Tulip Beds (Mount Stephen), which are both in Yoho National Park (British Columbia), quantitative assessments of the Burgess Shale have remained limited. Here we first provide a detailed quantitative overview of the diversity and structure of the Marble Canyon Burgess Shale locality based on 16,438 specimens. Located 40 km southeast of the Walcott Quarry in Kootenay National Park (British Columbia), Marble Canyon represents the youngest site of the “thick” Stephen Formation. We then combine paleoecological data sets from Marble Canyon, Walcott Quarry, Tulip Beds, and Raymond Quarry, which lies approximately 20 m directly above the Walcott Quarry, to yield a combined species abundance data set of 77,179 specimens encompassing 234 species-level taxa. Marble Canyon shows significant temporal changes in both taxonomic and ecological groups, suggesting periods of stasis followed by rapid turnover patterns at local and short temporal scales. At wider geographic and temporal scales, the different Burgess Shale sites occupy distinct areas in multivariate space. Overall, this suggests that the Burgess Shale paleocommunity is far patchier than previously thought and varies at both local and regional scales through the “thick” Stephen Formation. This underscores that our understanding of Cambrian diversity and ecological networks, particularly in early animal ecosystems, remains limited and highly dependent on new discoveries.

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.

The exceptionally preserved fossils entombed in the deposits of sediment-gravity flows in the Cambrian Burgess Shale of British Columbia have been fundamental for understanding the origin of major animal groups during the Cambrian explosion. More recently, they have been used to investigate the evolution of community structure; however, this assumes that the fossil assemblage represents an in-life community. Here we test this assumption for the first time based on experimental and field approaches. We use flume experiments to create analog flows and show that transport of the polychaete Alitta virens over tens of kilometers does not induce significantly more damage beyond that already experienced due to normal decay processes. Integration of experimental results with taphonomic assessment of fossils and sedimentological analysis suggests that the organisms of the Burgess Shale in the classic Walcott Quarry locality could have undergone substantial transport and may represent a conflation of more than one community. Exceptionally well preserved soft-bodied organisms could have been transported tens of kilometers by flows in the Burgess Shale and may not necessarily represent in-life communities, according to field observations and flume experiments on polychaetes.

International audience; Understanding the functioning of extinct ecosystems is a complicated knot of ecological, evolutionary, and preservational strands that must be untangled. For instance, anatomical and behavioral differences can profoundly alter fossilization pathways. This is particularly true in exceptionally preserved soft-bodied biotas that record the earliest phases of animal evolution during the Cambrian Explosion and the Ordovician Radiation. Herein, a novel method of data partitioning based on probabilistic modelling is developed to examine these processes for the Walcott Quarry, Burgess Shale, Canada (510Ma), and the Fezouata Shale, Morocco (c. 475Ma). The modelling shows that the mechanism for soft-tissue preservation in the Walcott Quarry is ecologically selective, favoring the endobenthos. This is not found in the Fezouata Shale. Taken in concert with bioturbation data, a new model of comparative preservation is developed based on sedimentary flow dynamics. This suggests that during the Cambrian 2 Explosion and Ordovician Radiation the most exceptional fossils sites must still be calibrated against each other to understand the unfolding evolutionary events and the ecological structuring of ancient animal communities.

The Tulip Beds locality on Mount Stephen (Yoho National Park, British Columbia) yields one of the most abundant and diverse (~10,000 specimens in 110 taxa) Burgess Shale fossil assemblages in the Canadian Rockies. Detailed semi quantitative and quantitative analyses of this assemblage suggest strong similarities with the Walcott Quarry on Fossil Ridge. Both assemblages are dominated by epibenthic, sessile, and suspension feeding taxa, mostly represented by arthropods and sponges and have comparable diversity patterns, despite sharing only about half the genera. However, the Tulip Beds has a higher relative abundance of suspension feeders and taxa of unknown affinity compared to the Walcott Quarry. These biotic variations are probably largely attributable to ecological and evolutionary differences between the two temporally distinct communities that adapted to similar, but not identical, environmental settings. For instance, the Tulip Beds is farther away from the Cathedral Escarpment than the Walcott Quarry. The Tulip Beds and Walcott Quarry assemblages are more similar to each other than either one is to the assemblages of the Chengjiang biota, although the relative diversity of major taxonomic groups and ecological patterns are similar in all assemblages. The conserved diversity patterns and ecological structures among sites suggest that the ecological composition of Cambrian Burgess Shale-type communities was relatively stable across wide geographic and temporal scales.

The degree to which the original community composition of the Middle Cambrian Burgess Shale was altered through transport and decay and how taphonomic conditions varied through time and across taxa is poorly understood. To address these issues, variation in fossil preservation was analyzed through a vertical succession of 26 bed assemblages, each representing a single obrution event, within the 7-m-thick Greater Phyllopod Bed of the Walcott Quarry. More than 50,000 specimens belonging to 158 genera—mostly benthic, monospecific and nonbiomineralized—were included in this analysis. The decay gradient of the polychaete Burgessochaeta setigera was used as a taphonomic threshold to estimate how far decay had proceeded in each bed assemblage. Qualitative comparisons of the degree of preservation of 15 species, representing an array of different body plans, demonstrate that all bed assemblages contain a mix of articulated and in situ dissociated or completely dissociated organisms interpreted respective...

In his work, Butterfield (1) makes several claims about carbonate cements in the Walcott Quarry Member of the Burgess Shale, including the contention that they are not regularly associated with fossil-bearing beds. This claim is inaccurate. Each millimeter- to centimeter-scale bed within the interval is cemented by calcite (figures 2–5 in ref. 2), including the exceptionally fossiliferous Great Marella Layer referred to in the work (1), which contains cements dispersed throughout its thickness and a prominently-cemented top (Fig. 1). The work by Butterfield (1) also suggests that the carbonate cements in the Burgess Shale were derived from recrystallization of coarse-grained carbonates and bioclasts during metamorphism. However, well-preserved skeletal fossils, such as Scenella, occur with clear margins and petrographic characteristics that are sharply distinct from the fine-grained carbonates that surround them, excluding the possibility that bioclasts were the original source of carbonate. Importantly, the cements in the Burgess Shale are entirely consistent in growth habit and distribution with the six other deposits analyzed in our study (3), including a nearby but less metamorphosed locality in the Stephen Formation, all of which clearly display early-displacive growth of carbonate at the seafloor. The assertion in the work by Butterfield (1) that seafloor precipitates are uncommon in the Cambrian is at odds with a large body of evidence (4).