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A columnal-bearing eocrinoid from the Cambrian Burgess Shale (British Columbia, Canada)

Published online by Cambridge University Press:  28 April 2015

Colin D. Sumrall
Affiliation:
Department of Earth and Planetary Sciences, University of Tennessee, Knoxville, TN 37996, USA, 〈csumrall@utk.edu〉
Samuel Zamora
Affiliation:
Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington DC, 20013–7012, USA

Abstract

A new eocrinoid ?Ubaghsicystis sp. from the middle Cambrian (Series 3, Stage 5) Burgess Shale is reported based on a single known specimen. This species extends the stratigraphic range of columnal-bearing eocrinoids in Laurentia significantly from Cambrian Stage 7 (Guzhangian) to Stage 5. It increases the diversity of echinoderms in this well-known fossil-Lagerstätte, provides the oldest evidence of columnal-bearing eocrinoids from Laurentia, and further documents the cosmopolitan distribution of middle Cambrian echinoderm clades.

Type
Articles
Copyright
Copyright © 2015, The Paleontological Society 

Introduction

The Burgess Shale of British Columbia, Canada, is the most famous Cambrian fossil locality in North America, and is often used to address questions about the timing and nature of the Cambrian Explosion and origin of animal phyla (Briggs et al., Reference Briggs, Erwin and Collier1994). Although excavations since the time of Walcott have uncovered an extraordinary diversity of animals (Collins, Reference Collins2009a), echinoderms remain scarce in collections and low in diversity when compared with similar-aged strata from Gondwana (Zamora, Reference Zamora2010; Smith et al., Reference Smith, Zamora and Álvaro2013).

Currently, only three unambiguous Burgess Shale echinoderms have been formally described. These include the plesiomorphic edrioasteroid Walcottidiscus typicalis Bassler, Reference Bassler1935 (Smith, Reference Smith1985; Zhao et al., Reference Zhao, Sumrall, Parsley and Peng2010), and the two eocrinoids Gogia stephenensis Sprinkle and Collins, Reference Sprinkle and Collins2006, and Lyracystis radiata Sprinkle, Reference Sprinkle1973 (Sprinkle and Collins, Reference Sprinkle and Collins2006). There are also poorly preserved ctenocystoids assignable to Ctenocystis that await formal description (Collins, Reference Collins2009b).

Other taxa have been considered by some to be echinoderms, including Echmatocrinus Sprinkle, Reference Sprinkle1973 (Sprinkle and Collins, Reference Sprinkle and Collins1998), and presumed holothuroids, including Eldonia ludwigi Walcott, Reference Walcott1911, Laggania cambria Walcott, Reference Walcott1911, Louisella pedunchlata Walcott, Reference Walcott1911, Mackenzia costalis Walcott, Reference Walcott1911, Redoubtia polypodia Walcott, Reference Walcott1911, and Portalia mira Walcott, Reference Walcott1911 (Walcott, Reference Walcott1918). Later authors have cast doubt on the echinoderm affinities of these fossils (Clarke, Reference Clarke1913; Madsen, Reference Madsen1957; Müller, Reference Müller1963; Ausich and Babcock, Reference Ausich and Babcock1998, Reference Ausich and Babcock2007; Daley et al., Reference Daley, Budd, Caron, Edgecombe and Collins2009).

The new eocrinoid from the Burgess Shale, while left under open nomenclature, has global implications for the timing of the origin of holomeric stems in eocrinoids. Prior to this discovery, the oldest Laurentian echinoderm bearing a holomeric stem was Eustipocystis minor Sprinkle, Reference Sprinkle1973, known from the middle Cambrian Guzhanguian. However, Gondwanan holomeric columnals predate those from Laurentia and are known from Cambrian Series 3 Stage 5 from several Gondwanan localities (Zamora et al., Reference Zamora, Clausen, Álvaro and Smith2010; Smith et al., Reference Smith, Zamora and Álvaro2013). This shows that the lack of early occurrences of holomeric stemmed echinoderms in Laurentia was a sampling artifact, and that by the middle Cambrian, such stems were present in both northern and southern continents.

The possible presence of Ubaghsicystis in the Burgess Shale provides further evidence for a cosmopolitan distribution of many echinoderm clades during the middle Cambrian. When Burgess Shale echinoderms are compared to nearly coeval deposits, it is noted that gogiids occur in Spain, France, Morocco, and China; Lyracystis occurs in China; Ubaghsicystis occurs in Morocco and Spain; Ctenocystis occurs in France and Australia; and Walcottidiscus (=Kailidiscus) occurs in China (Jell et al., Reference Jell, Burrett and Banks1985; Ubaghs, Reference Ubaghs1987; Parsley and Zhao, Reference Parsley and Zhao2006; Zamora, Reference Zamora2010; Sprinkle et al., Reference Sprinkle, Parsley, Zhao and Peng2011; Smith et al., Reference Smith, Zamora and Álvaro2013; Zamora et al., Reference Zamora, Lefebvre, Álvaro, Clausen, Elicki, Fatka, Jell, Kouchinsky, Lin, Nardin, Parsley, Rozhnov, Sprinkle, Sumrall, Vizcaïno and Smith2013). Compared with other North American localities, only two echinoderm clades (Soluta and Stylophora) are missing from the Burgess Shale (Ubaghs and Robison, Reference Ubaghs and Robison1985, Reference Ubaghs and Robison1988; Sumrall and Sprinkle, Reference Sumrall and Sprinkle1999). Further research on Burgess Shale faunas may yet discover specimens of these clades as shown by the fortuitous discovery of ?Ubaghsicystis sp.

Systematic paleontology

Subphylum Blastozoa Sprinkle, Reference Sprinkle1973

Class Eocrinoidea Jaekel, Reference Jaekel1918

Family uncertain

Genus Ubaghsicystis Gil Cid and Domínguez Alonso, Reference Gil Cid and Domínguez Alonso2002

Type species

Ubaghsicystis segurae Gil Cid and Domínguez Alonso, Reference Gil Cid and Domínguez Alonso2002 from the Cambrian of northern Spain.

Remarks

The new eocrinoid specimen from Burgess Shale is tentatively assigned to Ubaghsicystis based on the presence of a globular, polyplated theca bearing two feeding appendages and a slender, columnal-bearing stem. Although this taxon has a somewhat different size and distribution of thecal plates as compared to U. segurae, poor preservation precludes the description of a new species. Until now, Ubaghsicystis was only known from middle Cambrian deposits of Perigondwanan Spain and Morocco. The Spanish specimens were collected in the Cantabrian Mountains from the lower Languedocian section of the Oville Formation at the Barrios de Luna locality (León, North Spain). Specimens from Morocco are slightly older, and come from the Agzian Jbel Wawrmast Formation in its type locality (Gil Cid and Domínguez Alonso, Reference Gil Cid and Domínguez Alonso2002; Smith et al., Reference Smith, Zamora and Álvaro2013). The Agzian regional stage corresponds approximately with Cambrian Series 3, Stage 5, but the correlation of the Languedocian with global stages proposed by the International Subcommission on Cambrian Stratigraphy is problematic. Based on chemostratigraphic evidence, the lower Languedocian corresponds to Series 3 Stage 5 (see Álvaro et al., Reference Álvaro, Bauluz, Subías, Pierre and Vizcaïno2008). However, correlation based on trilobites and agnostids suggests it is correlative with the base of the Drumian (see Gozalo et al., Reference Gozalo, Chirivella Martorell, Esteve and Liñán2011).

?Ubaghsicystis sp.

Figure 1.1–1.3

Occurrence

BathyuriscusElrathina Zone, Cambrian Series 3, Stage 5. The specimen was collected in situ from a level 250 cm below the Phyllopod Bed in the Walcott Quarry, Mount Field, British Columbia.

Description

Theca of single known specimen globular, 7 mm in diameter, composed of polygonal plates, with at least 16–20 plates per exposed surface (Figs. 1, 2); specimen too poorly preserved to reveal exact disposition of plates or details of ornamentation, but plates appear slightly ridged and convex, some plates suggest small circular epispires along plate margins; two feedings appendages (brachioles) arise from oral spout (Figs. 1.1, 2); brachioles slender with massive, biserial brachiolar plates and very thin and small cover plates; connection with stem abrupt; on one side, two large plates at thecal base connect with the stem as in Ubaghsicystis segurae, no evidence seen for the position and plating of the periproct; stem relatively thin and gracile, holomeric with very short columnals (Figs. 1.2, 2), length unknown, but at least 10 mm visible; stem slightly wider close to theca and somewhat tapered, proximal width 0.8 mm, distalmost preserved diameter 0.3 mm. proximally with 6 columnals per mm.

Figure 1 ?Ubaghsicystis sp. showing a polyplated and globular theca, two brachioles, and a long stem constructed with low columnals. (1, 3) Part ROM 62443b and counterpart ROM 62443a immersed underwater with crossed polarized light, ×4.25; (2) Detail of proximal stem of ROM 62443a showing columnar construction, ×10.

Figure 2 Camera lucida drawing of Ubaghsicystis sp. ROM 62443a, showing two brachioles (b) arising from the thecal summit, a polyplated theca (t), and a gracile holomeric stem (s). The specimen is preserved overlapping an arthropod fragment (a), ×6.

Remarks

?Ubaghsicystis sp. differs from U. segurae by possessing a slightly less tumid theca and a more gracile stem. The poor preservation of the Burgess Shale specimen precludes formal description until better-preserved material is discovered.

The presence of ?Ubaghsicystis sp. in Laurentia is important because it provides new data on the timing of holomeric columnals in Laurentian eocrinoids. While this feature appears by the Cambrian Series 3 Stage 5 in several Gondwanan localities (Zamora et al., Reference Zamora, Clausen, Álvaro and Smith2010, Smith et al., Reference Smith, Zamora and Álvaro2013), the oldest previously reported Laurentian eocrinoid with holomeric columnals in the stem is the Guzhanguian Eustipocystis minor Sprinkle, Reference Sprinkle1973, thus extending the range of Laurentian holomeric columnals by 10 Myr. ?Ubaghsicystis sp. also adds another suspension-feeding echinoderm to the Burgess Shale assemblage.

Material

Part and counterpart (ROM 62443a, b), housed at the Royal Ontario Museum, Toronto, Ontario.

Acknowledgments

We acknowledge the support of the Royal Ontario Museum curatorial staff, especially J.-B. Caron and P. Fenton, and the Royal Ontario Museum Burgess Shale Project no. 29 for collecting the specimen described in this paper. S.Z. was funded by a postdoctoral fellowship at the Smithsonian Institution (Springer fund). Additional funding was provided by NSF DEB-1036260. J.A. Waters of Appalachian State University and B. Lefebvre of the Université Lyon provided valuable input to improve the manuscript.

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Figure 0

Figure 1 ?Ubaghsicystis sp. showing a polyplated and globular theca, two brachioles, and a long stem constructed with low columnals. (1, 3) Part ROM 62443b and counterpart ROM 62443a immersed underwater with crossed polarized light, ×4.25; (2) Detail of proximal stem of ROM 62443a showing columnar construction, ×10.

Figure 1

Figure 2 Camera lucida drawing of Ubaghsicystis sp. ROM 62443a, showing two brachioles (b) arising from the thecal summit, a polyplated theca (t), and a gracile holomeric stem (s). The specimen is preserved overlapping an arthropod fragment (a), ×6.