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SMALL SHELLY FOSSILS and CARBON ISOTOPES from the EARLY CAMBRIAN (STAGES 3–4) MURAL FORMATION of WESTERN LAURENTIA by CHRISTIAN B

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SMALL SHELLY FOSSILS and CARBON ISOTOPES from the EARLY CAMBRIAN (STAGES 3–4) MURAL FORMATION of WESTERN LAURENTIA by CHRISTIAN B [Papers in Palaeontology, 2020, pp. 1–33] SMALL SHELLY FOSSILS AND CARBON ISOTOPES FROM THE EARLY CAMBRIAN (STAGES 3–4) MURAL FORMATION OF WESTERN LAURENTIA by CHRISTIAN B. SKOVSTED1 , UWE BALTHASAR2 , JAKOB VINTHER3 and ERIK A. SPERLING4 1Department of Palaeobiology, Swedish Museum of Natural History, Box 50007, Stockholm, SE-104 05, Sweden; [email protected] 2School of Geography, Earth & Environmental Sciences, University of Plymouth, Plymouth, PL4 8AA, UK 3Schools of Biological Sciences & Earth Sciences, Life Sciences Building, University of Bristol, Bristol, BS8 1TQ, UK 4Department of Geological Sciences, Stanford University, Stanford, CA 94305, USA Typescript received 10 October 2019; accepted in revised form 23 January 2020 Abstract: The extraordinary window of phosphatized and species Hipponicharion perforata sp. nov. and Pseudobeyri- phosphatic small shelly fossils (SSF) during the early and chona taurata sp. nov. Compared with contemporaneous middle Cambrian is an important testament to the radiation faunas from western Laurentia, the fauna is relatively diverse, of biomineralizing metazoans. While SSF are well known particularly in taxa with originally phosphatic shells, which from most Cambrian palaeocontinents during this time appear to be associated with archaeocyathid build-ups. This interval, western Laurentia has relatively few SSF faunas. suggests that the generally low faunal diversity in western Here we describe a diverse SSF fauna from the early Cam- Laurentia may be at least partly a consequence of poor sam- brian (Stages 3–4) Mural Formation at three localities in pling of suitable archaeocyathan reef environments. In Alberta and British Columbia, Canada, complemented by addition, the tommotiid Canadiella filigrana appears to be of carbon isotope measurements to aid in a potential future biostratigraphical significance in Cambrian Stage 3 strata of bio-chemostratigraphic framework. The fauna expands the western Laurentia, and the unexpected high diversity of recorded SSF assemblage diversity in western Laurentia and bradoriid arthropods in the fauna also suggests that this includes several brachiopods, four bradoriids, three chancel- group may prove useful for biostratigraphical resolution in loriids, two hyoliths, a tommotiid and a helcionellid mollusc the region. as well as echinoderm ossicles and specimens of Microdic- tyon, Volborthella and Hyolithellus. New taxa include the Key words: small shelly fossils, Brachiopoda, Bradoriida, tommotiid genus Canadiella gen. nov., the new bradoriid biostratigraphy, carbon isotope stratigraphy, Cambrian. L OWER Cambrian fossils of the Mural Formation have carbonate-dominated lower Mural Formation and from been studied for over 100 years and a rich fauna of trilo- carbonate storm beds within the middle Mural bites and brachiopods has been documented (Walcott Formation. 1913; Fritz & Mountjoy 1975; Fritz 1992; Balthasar 2004, In the early Cambrian (Terreneuvian and Cambrian 2007, 2008, 2009). The discovery of exceptionally pre- Series 2), small shelly fossils (SSF) constitute a significant served fossils such as soft-shelled brachiopods (Balthasar portion of the total taxonomic diversity of the metazoan & Butterfield 2009), anomalocaridids and other non- fossil record (Maloof et al. 2010). SSF share nothing mineralized biota (Sperling et al. 2018) has also sparked beyond a commonly small size range (typically < 2 mm; interest in the formation. However, Burgess Shale-type which may be taphonomically biased for some taxa; Martı exceptional preservation in the Mural Formation is at best Mus et al. 2008) and the fact that the fossils are resistant marginal (a Tier 3 deposit), and there is little geochemical to digestion of the carbonate host rock in weak acids evidence for anoxic conditions that might result in more (fossils are typically either phosphatic by original compo- spectacular preservation (Sperling et al. 2018). The bulk sition or secondarily phosphatized or silicified calcareous of the known fauna has been derived from mudstones of shells). The fossils include various shells, sclerites, spicules the middle part of the Mural Formation. Here, we and other skeletal elements belonging to a multitude of describe skeletal fossils from both the underlying early animal groups representing stem and crown © 2020 Authors. doi: 10.1002/spp2.1313 1 Papers in Palaeontology published by John Wiley & Sons Ltd on behalf of The Palaeontological Association. This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited. 2 PAPERS IN PALAEONTOLOGY members of various phyla across the metazoan tree of life The excellent preservation and relatively high taxo- (Budd & Jensen 2000; Kouchinsky et al. 2012). nomic diversity presented here makes the fauna of the Relatively few SSF faunas have been described from Mural Formation one of the most diverse and best pre- the Cambrian of western Laurentia. From the earliest served SSF faunas described from the lower Cambrian of Cambrian (Fortunian), Conway Morris & Fritz (1980) western Laurentia. The results highlight that an increased reported a single protoconodont specimen (probably sampling focus on archaeocyathan reefs and associated from the Ingta Formation in the Mackenzie Mountains sediments might increase the known SSF diversity in of north-western Canada; see Aitken 1989), and Pyle western Laurentia. Our results suggest that specific SSF et al. (2006) described a fauna including possible animal taxa such as tommotiids and bradoriid arthropods have a embryos, anabaritids and problematic fossils from the good potential for biostratigraphical resolution in Cam- Wernecke Mountains of Yukon, Canada. However, SSF brian Stage 3 of western Laurentia, although further work records from Laurentia are more common in the late is required to realize this potential. With the combination early Cambrian (Series 2), particularly along the present of carbon isotope chemostratigraphic data in conjunction eastern margin of North America. In western Laurentia, with SSF data, as we present here, it may be possible in published Series 2 accounts from the Great Basin of the the future to build an improved bio-chemostratigraphic western USA are limited to brachiopods and a handful framework. Ultimately this will be key to placing western of generally poorly preserved taxa (Signor & Mount Laurentian SSF diversity in the global picture and deter- 1986; Skovsted 2006a; Skovsted & Holmer 2006; Wotte mining if the observed low diversity is due to sampling, & Sundberg 2017; Pruss et al. 2019). The same applies ecology, or taphonomic effects. to the fauna from the Sekwi Formation of the Macken- zie Mountains of north-western Canada (Voronova et al. 1987) and Sonora, Mexico (McMenamin 1984; Devaere GEOLOGICAL SETTING et al. 2019). In addition, the occurrence of single prob- lematic SSF have been reported from different regions The Mural Formation is located in the southern Canadian of western North America, i.e. Salterella (Fritz & Cordillera and was deposited on the western Laurentian Yochelson 1988), Volborthella (Hagadorn & Waggoner margin, approximately during the rift–drift transition and 2002), Microdictyon (Bengtson et al. 1986), and ‘Lap- initial Palaeozoic flooding of North America (Sauk trans- worthella’ filigrana (Conway Morris & Fritz 1984). gression) (Pope et al. 2012). The Mural was deposited in Hence, the SSF fossil record from western Laurentia is a relatively shallow-water environment, with the Lauren- relatively meagre compared with contemporaneous fau- tian craton to the east and deeper water conditions devel- nas from eastern Laurentia (Skovsted 2006b; Skovsted & oping to the west. The Mural Formation itself thins Peel 2007, 2011) and other palaeocontinents (Qian & substantially across the Peace River Arch, a major east- Bengtson 1989; Bengtson et al. 1990; Kouchinsky et al. north–east-trending structure in northern Alberta and 2012, 2015). British Columbia (Fig. 1; McMechan 1990). However, the Biostratigraphic subdivision and correlation of Cam- tripartite stratigraphic motif of the Mural is recognized brian strata traditionally rests on trilobites (Shergold & from Mexico to Yukon during the Lower Cambrian (Ser- Geyer 2003; Zhu et al. 2019). However, a significant part ies 2, Age 3 and 4; Waucoban Series, Montezuman– of the Cambrian Period is pre-trilobitic (Terreneuvian) Dyeran Stages in a North American timescale; Nevadella– and in the overlying unnamed Cambrian Series 2, trilo- Bonnia-Olenellus trilobite zones in older literature). This bite faunas have proven to be highly endemic with result- motif consists of a lower carbonate (often archaeocyathan ing problems for intercontinental correlation. Carbon limestone mounds/biostromes and ooid grainstone), a isotope stratigraphy has also emerged as a robust correla- medial shale/siltstone, and an upper carbonate, again tion method in lower Cambrian strata (e.g. Smith et al. often containing archaeocyaths (Pope et al. 2012). In 2016) but both patterns of negative/positive excursions most localities, the Nevadella–Bonnia-Olenellus boundary, and absolute values can be non-unique. Consequently, which is also the Montezuman–Dyeran boundary, is additional sources of age control are needed. Recently, located in the medial shale (Fig. 2). Additional data on non-trilobite shelly fossils were used to define a new bios- the geology, sedimentology, and palaeontology
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