Assemblage: Thirteen More Short Stories

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Liangshanella Fig. Importantly, two arthropod taxa previously assumed to be endemic to the early Cambrian Chengjiang biota, the problematic Primicaris 16 Fig. The presence of these taxa within the Marble Canyon assemblage significantly expands their geographical and temporal ranges, and indicates that their absence from younger Burgess Shale-type assemblages was not the result of endemism or extinction, but was controlled by ecological and taphonomic factors.

This finding, together with those from the recently described Early Ordovician Fezouata biota 17 , suggests that the longevity and geographical ranges of soft-bodied Cambrian taxa at both high and low taxonomic ranks may be severely underestimated. Among the new taxa recovered from the Marble Canyon locality, arthropods are prominent in both diversity and abundance, and include at least 12 new soft-bodied morphotypes. These new forms will aid in understanding the relationships and disparity of megacheirans 18 , 19 , 20 , 21 and bivalved arthropods 22 , 23 , two groups that continue to play a pivotal role in debates surrounding the early evolution of arthropods.

The new evidence emphasizes the predatorial function of leanchoiliid appendages and the likely connections between the dinocaridid and bivalved bodyplans, which have only tentatively been explored so far 24 , but are of critical importance to the evaluation of overall arthropod disparity.

Another new bivalved arthropod Fig. This confirms that an anterior, post-oral, chelate appendage appeared early, possibly deep within the stem-group of arthropods.

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The assemblage also yields the first Mollisonia 25 specimen to reveal extensive details of cephalic appendages, together with eyes showing retinal and brain tissues, including putative neuropils Fig. Mollisonia was previously known only from elements of the exoskeleton 10 , with soft tissues never before observed.

The new material suggests that the neglected taxon Houghtonites 26 actually belongs to Mollisonia , and that this animal was an active benthic predator, based on the presence of large lateral eyes and long cephalic appendages.

The consistently high fidelity of preservation in the Marble Canyon material is striking, and expands the range of internal tissues preserved in Cambrian Burgess Shale-type fossils. These include a putative liver and heart in Metaspriggina Fig. We also report for the first time the preservation of neural tissues in Burgess Shale material Fig. Additional new anatomical features include a putative gut and indeterminate organs in the enigmatic lophotrochozoan hyolith Haplophrentis Fig.

Appendages in the trilobite Kootenia Supplementary Fig. To date, collection efforts at Marble Canyon have been limited to a single field season of 15 days. Yet, remarkably, the rarefaction curve for Marble Canyon arthropods is markedly steeper than those for arthropods of the Walcott Quarry and Chengjiang assemblages, or for the closely adjacent Stanley Glacier assemblage 10 Fig.

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Differences in diversity and composition Fig. The density of fossils in the Marble Canyon assemblage, normalized to volume of rock excavated, is significantly higher than that of the Chengjiang or Stanley Glacier localities, and is comparable only to the Walcott Quarry Fig. Parallel trends in fossil density and diversity indicate that additional collection at Marble Canyon holds great promise for the recovery of new taxa and for enhancing the present understanding of the taxonomic diversity and morphological disparity of the Cambrian fauna.

Black bars indicate interquartile range of log density; white dots indicate median of log density; surrounding coloured areas indicate Kernel density plots; MC, Marble Canyon magenta ; MF, Mafang black , WQ, Walcott Quarry blue. Patterns in fossil associations point to the recurrence of a similar community-type across the fossil-rich interval Supplementary Data 1.

While this suggests that relatively consistent taphonomic, ecological and environmental conditions prevailed throughout the studied interval, it also highlights important differences in content between Marble Canyon and other Cambrian soft-bodied assemblages elsewhere. Sessile organisms, in particular sponges and brachiopods, which are especially abundant and diverse at the Walcott Quarry 5 and Chengjiang 28 and to a lesser extent at Stanley Glacier 10 , are consistently rare at Marble Canyon.

Differences in taxonomic composition, as well as in diversity and density, emphasize the role of palaeoenvironment in regulating ecological and taphonomic controls that determined the composition of soft-bodied fossil assemblages, even across small spatial scales. While sampling methodologies, accessibility and collecting efforts vary greatly among Burgess Shale-type deposits, this finding confirms that extraordinarily fossil-rich intervals of the Burgess Shale are neither limited to the type area nor to the lower and middle parts of the formation, which have historically been the focus of investigation.

This discovery suggests that the potential of the Burgess Shale to yield exceptional biotas across a spectrum of ecological, environmental and taphonomic gradients remains largely untapped.


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The particularly large proportion of new taxa that occur within the Marble Canyon assemblage, which lies in close temporal and geographical proximity to the best sampled Cambrian soft-bodied assemblage, the Walcott Quarry, suggests that the present understanding of the diversity and disparity of the earliest complex ecosystems of the Phanerozoic remains in its infancy, and awaits future discoveries.

All fossil localities visited during the research are protected under the National Parks Act and all fossils and geological samples were collected under a Parks Canada collection and research permit YNP Shelly fossils from carbonate horizons were collected in the lab via acid maceration dilute acetic and formic acids.


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All available exposures of the Stephen Formation in the vicinity of Marble Canyon were measured, sampled and described in cm-scale detail in the field. Observations of sections on Mount Whymper were included in producing a generalized composite section Fig. Where extensive talus aprons were found to cover portions of the Stephen Formation, a laser rangefinder was used to determine the stratigraphic thickness of the covered interval. Samples were analysed in thin section, polished slab and X-radiograph. All fossils were identified to the lowest possible taxonomic rank usually species and some specimens were prepared using a micro-engraving tool equipped with a tungsten bit.

EstimateS 9. Density estimates were calculated for intervals with more than specimens for all phyla and more than 32 specimens for arthropods only; Supplementary Data 2 ; Supplementary Note 2. Kruskall—Wallis tests were significant for all comparisons of density among the Mafang, Marble Canyon and Walcott Quarry assemblages.

For the Walcott Quarry and Marble Canyon localities, P values of Mann—Whitney tests for specimens and taxa in the whole assemblage were 0. When limiting the tests to arthropods, the P value was insignificant for specimen counts 0. Similar tests were run using the data from the Stanley Glacier assemblage; although the number of specimens overall was too low to compare with the others, density estimates were found to be close to that of the Mafang assemblage. How to cite this article: Caron, J. A new phyllopod bed-like assemblage from the Burgess Shale of the Canadian Rockies.

Conway Morris, S. Burgess Shale faunas and the Cambrian explosion. Science , — Whittington, H. The Burgess Shale: history of research and preservation of fossils. Convention Part 1 Extraordinary Fossils : — The community structure of the Middle Cambrian phyllopod bed Burgess Shale. Palaeontology 29 , — Caron, J. Ou, Q. Cambrian lobopodians and extant onychophorans provide new insights into early cephalization in Panarthropoda.

Zhang, W. Preliminary notes on the occurrence of the unusual trilobite Naraoia in Asia. Acta Paleontol. Sinica 24 , — Collins, D. New Burgess Shale fossil sites reveal Middle Cambrian faunal complex. Geology 38 , — Gaines, R. Geology 40 , — The paleoredox setting of Burgess Shale-type deposits. The Burgess Shale animal Oesia is not a chaetognath: a reply to Szaniawski Acta Palaeontol. Zhang, X.

Cambrian naraoiids Arthropoda : morphology, ontogeny, systematics, and evolutionary relationships. Van Roy, P. Ordovician faunas of Burgess Shale type. Nature , — Hou, X. Arthropods of the Lower Cambrian Chengjiang fauna, southwest China.


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  4. Fossils and Strata 45 , — The oldest arthropods and the origin of Crustacea. Acta Zool. Haug, J. Morphology and function in the Cambrian Burgess Shale megacheiran arthropod Leanchoilia superlata and the application of a descriptive matrix. BMC Evol. Cotton, T. The phylogeny of arachnomorph arthropods and the origin of the Chelicerata.

    Earth Sci. Briggs, D. The early radiation and relationships of the major arthropod groups. Legg, D. Cambrian bivalved arthropod reveals origin of arthrodization. Although the states remained sovereign and independent, no state was to impose restrictions on the trade or the movement of citizens of another state not imposed on its own. Movement across state lines was not to be restricted. To amend the Articles, the legislatures of all thirteen states would have to agree.

    This provision, like many in the Articles, indicated that powerful provincial loyalties and suspicions of central authority persisted. In the s—the so-called Critical Period—state actions powerfully affected politics and economic life. For the most part, business prospered and the economy grew. Expansion into the West proceeded and population increased. National problems persisted, however, as American merchants were barred from the British West Indies and the British army continued to hold posts in the Old Northwest, which was named American territory under the Treaty of Paris.

    Notes on the Assemblage

    These circumstances contributed to a sense that constitutional revision was imperative. Still, national feeling grew slowly in the s, although major efforts to amend the Articles in order to give Congress the power to tax failed in and The year after the failure of , the Constitutional Convention met in Philadelphia and effectively closed the history of government under the Articles of Confederation. To all to whom these Presents shall come, we the undersigned Delegates of the States affixed to our Names send greeting.

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    And that the articles thereof shall be inviolably observed by the states we respectively represent, and that the union shall be perpetual. In Witness whereof we have hereunto set our hands in Congress. Done at Philadelphia in the state of Pennsylvania the ninth day of July in the Year of our Lord one Thousand seven Hundred and Seventy-eight, and in the third year of the independence of America. Start your free trial today. But if you see something that doesn't look right, click here to contact us!

    Subscribe for fascinating stories connecting the past to the present. From to , the Continental Congress served as the government of the 13 American colonies and later the United States. The First Continental Congress, which was comprised of delegates from the colonies, met in in reaction to the Coercive Acts, a series of measures