SEPTEMBER 2010

■
VOL. 38 NO. 9

ISSN 0091-7613 SEPTEMBER 2010 ■ VOL. 38 NO. 9 ■ P. 769–864

INSIDE:
Earthquakes Get Hot under the Collar, p. 771
Antarctic DJ for Tropical Dance, p. 783
The Himalaya Show Their Age, p. 807
Rodrigues Kicks the Bucket, p. 855
 Downloaded from geology.gsapubs.org on September 4, 2010

A new Burgess Shale–type assemblage from the “thin” Stephen

Formation of the southern Canadian Rockies
Jean-Bernard Caron1, Robert R. Gaines2, M. Gabriela Mángano3, Michael Streng4, and Allison C. Daley4
1
Department of Natural History-Palaeobiology, Royal Ontario Museum, 100 Queen’s Park, Toronto, Ontario M5S 2C6, Canada
2
Pomona College, Geology Department, 185 E. Sixth Street, Claremont, California 91711, USA
3
Department of Geological Sciences, University of Saskatchewan, 114 Science Place, Saskatoon, Saskatchewan S7N 5E2, Canada
4
Department of Earth Sciences, Uppsala University, Villavägen 16, 75236 Uppsala, Sweden

ABSTRACT
51°39′N
A new Burgess Shale–type assemblage, from the Stephen Forma-

AL
116°53′W

BE
tion of the southern Canadian Rocky Mountains, is described herein.
YOHO 1000 km

RT
It occurs near Stanley Glacier in Kootenay National Park, 40 km
N.P. CANADA

A
southeast of the type area near Field, British Columbia. While at least
a dozen Burgess Shale localities are known from the “thick” Stephen CaCtahth
eed
Formation, the Stanley Glacier locality represents the first discovery F

drar
Field F EEs

lal
of Burgess Shale–type fossils from the “thin” Stephen Formation.
F

scca
The Cathedral Escarpment, an important regional paleotopographic

ra
F

m p
rpenm
feature, has been considered important to the paleoecologic setting F

t
F

en
and the preservation of the Burgess Shale biota. However, the Stanley

t
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 anom- KOOTENAY Banff
alocaridid Stanleycaris hirpex n. gen., n. sp. (new genus, new spe-
N N.P.
BR OL
cies). Nektonic or nektobenthic predators represent the most diverse F 1

Stanley
C
IT UM
Monarch

115°30′
IS B
group, whereas in relative abundance, the assemblage is dominated
by typical Cambrian shelly benthic taxa. The low diversity of both
H IA Glacier Cirque
the benthic taxa and the ichnofauna, which includes diminutive trace F
fossils associated with carapaces of soft-bodied arthropods, suggests 10 km 51°0′
a paleoenvironment with restrictive conditions. The Stanley Glacier
assemblage expands the temporal and geographic range of the Bur- Figure 1. Stanley Glacier fossil locality (for close-up, see Fig. DR1
gess Shale biota in the southern Canadian Rockies, and suggests that [see footnote 1]) in relation to Burgess Shale–type localities (F) in
“thick” Stephen Formation (dark gray areas). Modified from Collins
Burgess Shale–type assemblages may be common in the “thin” Ste- et al. (1983) and Stewart (1991).
phen Formation, which is regionally widespread.

INTRODUCTION
Cambrian fossil Lagerstätten provide unique insights into the early
diversification of the Metazoa in the aftermath of the Cambrian explo-
sion (Conway Morris, 1992). Among these deposits, the middle Cambrian
(Series 3) Burgess Shale of British Columbia, Canada, is certainly the best
known. The Burgess Shale biota occurs at several localities, from the type
area in Yoho National Park near Field, British Columbia, to ~60 km south-
east, near Monarch Cirque (Fig. 1). At each of these localities, Burgess
Shale–type (BST) fossils occur within the thick expression of the Stephen
Formation (Aitken, 1981; Collins et al., 1983; Stewart, 1991). This suc-
cession of mudstones and carbonates (~300 m), also referred to as the Bur-
gess Shale Formation (Fletcher and Collins, 1998), accumulated offshore
of a large submarine cliff, termed the Cathedral Escarpment. Because of
the direct association of fossil localities with the Cathedral Escarpment,
many have considered the escarpment important to the ecological setting
or preservation of the Burgess Shale biota (e.g., Conway Morris, 1986).
However, questions remain regarding the temporal and geographic dis-
tribution of the Burgess Shale biota and the range of paleoenvironmental
conditions in which BST fossils occur. In particular, a thinner (typically
<60 m) succession of carbonates and mudstones termed the “thin” Ste-
phen Formation, representing deposition shoreward of the escarpment,
has not been well explored for BST fossils despite its association with the
thicker succession.
Here we describe a new BST assemblage from the “thin” Stephen
Formation at Stanley Glacier, Kootenay National Park, ~40 km southeast
of the type area (Fig. 1; see Fig. DR1 in the GSA Data Repository1). This
assemblage was excavated by a Royal Ontario Museum party in 2008 fol-
lowing recent reports of BST fossils collected from talus material (Caron
et al., 2010; Daley et al., 2009; Rigby and Collins, 2004; Vannier et al.,
2007). The Stanley Glacier BST assemblage expands the diversity and
ecological range of the Burgess Shale biota, widens its known geographi-
cal and paleoenvironmental range, and fills a temporal gap in the Cam-
brian stratigraphic record of BST deposits.
GEOLOGICAL SETTING
The Stephen Formation in the Stanley Glacier area conformably
overlies cryptalgal dolostones of the Cathedral Formation, rather than its
deep-water equivalent, the Takkakaw Tongue, supporting an assignment
to the “thin” Stephen. In the study area, the Stephen Formation is 32.8 m
thick (Fig. 2), well within the range of “thin” Stephen thicknesses known
regionally (16–164 m) and outside of the range of “thick” Stephen sec-
tions (276–370 m) in the region (Aitken, 1997). No evidence of an escarp-
ment has been reported previously from the Stanley Glacier area or identi-
fied in reconnaissance study of the region.
1
GSA Data Repository item 2010228, Figures DR1–DR6, Table DR1
(specimens counts and relative abundance chart), and Item DR1 (description of
Stanleycaris hirpex), is available online at www.geosociety.org/pubs/ft2010.htm,
or on request from editing@geosociety.org or Documents Secretary, GSA, P.O.
Box 9140, Boulder, CO 80301, USA.

© 2010 Geological Society of America. For permission to copy, contact Copyright Permissions, GSA, or editing@geosociety.org.
GEOLOGY,
Geology, September
September 2010
2010; v. 38; no. 9; p. 811–814; doi: 10.1130/G31080.1; 3 figures; Data Repository item 2010228. 811
 Downloaded from geology.gsapubs.org on September 4, 2010

bottom-flowing currents is evidenced by randomly oriented clay micro-

fabric (O’Brien et al., 1980), and by the occurrence of common skeletal
lags and rare tool marks at bed bases. Event beds are amalgamated with
little to no intervening background sediment and range from 1 to 11 mm
in thickness (Fig. DR2).

BURGESS SHALE–TYPE ASSEMBLAGE

The BST assemblage (24 taxa) at Stanley Glacier occurs in the clay-
stones of cycle 5, from 27.01 to 30.02 m above the base of the “thin”
Stephen Formation (Fig. 2). Fossils (N = 1098) were collected or counted
through systematic excavations of 23 intervals (6–22 cm in thickness,
average 10 cm) representing a complete section of cycle 5. Although some
variation is present, the composition of each interval is comparable and
dominated by shelly organisms (Table DR1). Fossils were also collected
from talus at several outcrops (N = 687), over an area spanning at least
10 km2 (Fig. DR1).
The shelly component of the Stanley Glacier assemblage includes
typical Cambrian taxa: hyolithids, brachiopods, and trilobites. All three
groups are also present in carbonate event beds of the Stephen Formation
with some shared genera (e.g., Spencella and Lingulella), but with few
species in common (Fig. DR3). BST fossils occur in claystones, where
soft-bodied arthropods represent the most diverse taxonomic group (9
taxa, 39% of the species), but compose only 11.8% of specimens. The
top five taxa in rank order of abundance account for 84.6% of specimens
(Table DR1) and include the enigmatic mollusc-like Haplophrentis cari-
natus (Fig. 3A; Fig. DR5C), brachiopods (Lingulella waptaensis and an
indeterminate acrotretid), and “ptychopariid” trilobites (Spencella sp.
and Ehmaniella burgessensis; Figs. DR4E and DR4F). The next four

Figure 2. Stratigraphic section of the “thin” Stephen Formation

near Stanley Glacier, including upper 6 m of Cathedral Formation
and lower 5 m of Eldon Formation. 1–6—lithostratigraphic cycles,
interpreted as parasequences; BST—occurrence of Burgess Shale–
type assemblage in claystones of cycle 5. M—mudstone; W—wacke-
stone; P—packstone; Sh—shale; Si—siltstone; SS—sandstone.

Although the “thin” Stephen Formation has been considered to rep-

resent deposition under shallow-water conditions (e.g., Fletcher and Col-
lins, 1998), the absence of hummocky cross-stratification indicates that
the entire Stephen Formation at Stanley Glacier was deposited below
storm wave base. No grading, scour, or cross-bedding is present within
the entire thickness of the Stephen Formation.
The shale-dominated succession is organized into six shale-wacke-
stone cycles, interpreted as parasequences (Fig. 2). Similar cycles were
documented in the study area by Aitken (1997), who provided regional-
scale evidence that they represent upward shallowing, expressed as
increasing carbonate input relative to shale. At Stanley Glacier, cycle
bases are abrupt and marked by the appearance of monotonous, calcare-
ous claystones that have millimeter-scale laminae. Near cycle tops, nodu-
lar carbonates appear within claystones and pass upward into thin-bedded
nodular wackestones. Storm-generated packstones (12–32 cm) bearing
oncoids and other platform-derived bioclasts deposited as single events Figure 3. Burgess Shale–type (BST) fossils from the “thin” Stephen
occur most frequently near cycle tops. Formation, Stanley Glacier area. A: Haplophrentis carinatus, ROM
(Royal Ontario Museum) 59943. B: Isolated appendages of Stan-
The top of the “thin” Stephen Formation is marked by the top of leycaris hirpex n. gen., n. sp. (left; see also Data Repository Item
the highest shale in the succession (Aitken, 1997), and the transition to DR1B [see footnote 1]) and Hurdia victoria, ROM 59944. C: Sidneyia
the thin-bedded nodular wackestones of the overlying Eldon Formation inexpectans, ROM 59945. D: Diagoniella cyathiformis, ROM 59946.
is gradational. Like other BST deposits (e.g., Gaines et al., 2005), the E: Isolated appendages of Anomalocaris canadensis, ROM 59947.
F: Great appendage arthropod A, part, frontal appendage indicated
claystones bearing the BST assemblage are ultrafine grained (<20 µm; by arrow, ROM 59948 (see counterpart in Fig. DR4D). G: Mollisonid A
Fig. DR2) and are composed exclusively of clays with pervasive authi- arthropod, ROM 59949. H: Sipunculan A worm, ROM 59950 (see also
genic calcite cements (2–37 wt%). Event-driven deposition of clays from Figs. DR4G and DR4H). Scale bars = 1 cm.

812 GEOLOGY, September 2010

 Downloaded from geology.gsapubs.org on September 4, 2010
most abundant taxa are nektobenthic or nektonic soft-bodied arthropods
(Hurdia victoria, Stanleycaris hirpex n. gen., n. sp., Sidneyia inexpectans,
Figs. 3B and 3C; Tuzoia retifera, Item DR1, Figs. DR4A–DR4C, DR5A,
and DR5B), followed by the hexactinellid sponge Diagoniella cyathifor-
mis (Fig. 3D). The morphology of Hurdia and Sidneyia matches that seen
in the type localities, with Hurdia having morph B frontal appendages,
broad H elements, and highly toothed mouthparts (Fig. 3B; Fig. DR5B)
(Briggs, 1979; Daley et al., 2009). Stanleycaris hirpex n. gen. n. sp. is a
new anomalocaridid consisting of appendages with 11 podomeres bear-
ing 5 elongated, spiniferous ventral spines and 9 large, double-pointed
dorsal spines (Fig. 3B; Item DR1). These appendages are often paired
and associated with a peytoia-type mouthpart. One specimen of Tuzoia
is preserved with a pair of eyes and the first appendages known from this
bivalved arthropod, including two robust anterior appendages with at least
six podomeres each (Figs. DR4A–DR4C). Allonnia sp., the hexactinellid
sponge Hintzespongia bilamina, the alga Margaretia dorus, and the nek-
tonic arthropods Anomalocaris canadensis (Fig. 3E) and Isoxys carinatus
are rare. At least two other new arthropods are present in the assemblage.
Great appendage arthropod A is broadly similar to Fuxianhuia from the
Chengjiang biota (Hou, 1987) and has a pair of short frontal appendages,
a truncated head shield bearing one pair of large pedunculate eyes, and
at least 32 trunk somites associated with presumably biramous limbs
(Fig. 3F; Fig. DR4D). Another arthropod, mollisonid A, resembles Mol-
lisonia (Briggs et al., 2008), but has nine trunk segments (Fig. 3G). A
new putative sipunculan worm, sipunculan A (Fig. 3H; Figs. DR4G, and
DR4H), reminiscent of Chinese forms (Huang et al., 2004), is also present
(D.Y. Huang, 2009, personal commun.). The assemblage also includes a
tentaculate organism, Herpetogaster collinsi (Caron et al., 2010), and six
other indeterminate forms (Table DR1A), including a paterinid brachio-
pod with preservation of setae (Fig. DR4I).
TRACE FOSSILS AND TAPHONOMY
Ichnofossils are abundant on some bedding planes (Fig. DR6).
However, ichnodiversity of individual assemblages is relatively low (2–5
ichnotaxa) and biogenic structures are only shallowly penetrative with
weakly developed ichnofabrics. Trace fossils are typically millimeters
to micrometers in size, recording the activities of small in situ benthic
macrofauna and meiofauna including grazers, deposit feeders, possible
carnivores, and gardeners (agrichnia). These include simple trails (e.g.,
Gordia), bilobate trails (e.g., Cruziana problematica), shallow rosary bur-
rows (“Hormosiroidea”-like structures), and arthropod trackways (e.g.,
Diplichnites). Diminutive trails (typically <1.5 mm wide), similar to forms
described from Chengjiang (Zhang et al., 2007), are commonly associated
with carapaces of Hurdia, Sidneyia, and Tuzoia. The most abundant struc-
tures are simple and straight or curved to irregularly or regularly sinuous
trails that do not self-overcross (e.g., Helminthoidichnites, Helminthopsis,
and Cochlichnus). Tiny microburrows of deposit feeders (Alcynidiopsis)
are also present in direct association with carapaces.
Additional evidence of an in situ benthic assemblage comes from
body fossils. The presence of both isolated parts and articulated specimens
of the same species suggests minimal time-averaging assemblage with
limited transport (Kidwell and Flessa, 1995), as well as limited decay and
rapid burial (Allison, 1986). In addition, a worm preserved at the end of a
possible mucus tube or burrow trace (Fig. 3H; Figs. DR4G, and DR4H),
complete specimens of the sponge Diagoniella (Fig. 3D), probable molt
assemblages of Hurdia and Sidneyia, and clusters of articulated specimens
of Haplophrentis (Figs. DR5A–DR5C) with no evidence of size sorting
suggest in situ burial. However, this does not preclude limited transport or
delivery of nektonic animals from the water column (e.g., Anomalocaris
appendages, Tuzoia valves, Hurdia carapaces).
Scanning electron microscope–energy dispersive X-ray analysis
elemental mapping indicates that the remains of soft tissues are pre-
GEOLOGY, September 2010
served as carbonaceous compression fossils, as in other BST deposits
(Gaines et al., 2008). The “thin” Stephen Formation at Stanley Glacier
has undergone far less metamorphism than at the classic Burgess Shale
localities (Butterfield et al., 2007; Powell, 2003), as evidenced by the
absence of muscovite and dominance of illite over chlorite in X-ray dif-
fraction spectra of clay separates.
PALEOENVIRONMENTAL SYNTHESIS AND AGE
The “thin” Stephen Formation at Stanley Glacier was deposited in
a distal ramp setting in a region where no evidence of an escarpment is
present. This depositional setting is more similar to that of BST deposits
of western Laurentia (Brett et al. 2009; Elrick and Snider, 2002; Gaines
et al., 2005; Liddell et al., 1997; Rees, 1986; Webster et al., 2008) and
south China (Zhao et al., 2005; Zhu et al., 2001) than it is to the classic
Burgess Shale localities near Field (Fletcher and Collins, 1998; Gabbott et
al., 2008). The presence of complete specimens of hexactinellid sponges
indicates low background sedimentation rates, suggestive of a relatively
deep water environment (Carrera and Botting, 2008), as corroborated
by sedimentological evidence. The presence of the alga Margaretia and
arthropods with eyes, such as Hurdia and Sidneyia, suggests a living envi-
ronment within the photic zone. The different species composition of the
carbonate event beds presumably reflects the shallower water environ-
ments upslope from which they were derived.
The trilobites Ehmaniella burgessensis and Spencella sp. and strati-
graphic evidence presented here and in Aitken (1997) suggest that the
“thin” Stephen Formation at Stanley Glacier is within the upper part of
the Bathyuriscus-Elrathina Zone, probably in the Ehmaniella Subzone
(Sundberg, 1994). The BST biota is stratigraphically above the Burgess
Shale localities near Field, and below the BST biota of the Wheeler For-
mation (Utah, USA) (Conway Morris, 1992).
COMPARISONS WITH COEVAL DEPOSITS FROM THE
“THICK” STEPHEN AND SIGNIFICANCE OF THE STANLEY
GLACIER BURGESS SHALE–TYPE ASSEMBLAGE
About half of the genera from Stanley Glacier are also present at the
type localities on Fossil Ridge (Caron and Jackson, 2008; Conway Mor-
ris, 1986), but the overall low species diversity of soft-bodied forms and
the disproportionate abundance of shelly elements at Stanley Glacier is
more similar to other BST deposits of western Laurentia (Liddell et al.,
1997; Robison, 1991). The presence of soft-bodied predators or scaven-
gers occupying a diversity of ecological niches suggests a complex food
web, similar to those described from the type localities and the Cheng-
jiang biota (Caron and Jackson, 2008; Conway Morris, 1986; Dunne et al.,
2008). However, the great diversity of nektonic predators or scavengers,
represented mostly by soft-bodied arthropods, compared to the low diver-
sity of benthic organisms, is curious, suggesting a possible taphonomic
bias. Paleoenvironmental factors such as oxygen stress might also have
precluded the development of a species-rich assemblage (e.g., Peters,
2007), with the seafloor colonized only periodically by more cosmopoli-
tan or generalist taxa such as Haplophrentis or small ichnofauna.
Like other BST deposits of Laurentia, the Stanley Glacier assem-
blage occurs across a broad area (Fig. DR1) without direct association
with an escarpment. Considering the significant taxonomic overlap
between the Stanley Glacier and other fossil assemblages in the type area,
and evidence that an in situ benthos was present at least periodically (Alli-
son and Brett, 1995), the presence of high species diversity in the latter
assemblages reinforces the view that deposits adjacent to the Cathedral
Escarpment represent a unique paleoenvironmental setting.
Owing to the discovery of several new species within a small geo-
graphic area during this preliminary phase of exploration, we consider
it likely that future exploration and study will continue to yield new taxa
from the “thin” Stephen Formation, which is exposed over a broader area
813
 Downloaded from geology.gsapubs.org on September 4, 2010
regionally than the “thick” Stephen Formation (Aitken, 1997; Stewart,
1991), and will increase our understanding of the geographic range and
evolutionary significance of the Burgess Shale biota.
ACKNOWLEDGMENTS
We thank Nigel Hughes and Peter Allison for their constructive reviews,
David Rudkin for comments on trilobites, and Parks Canada for a Research and
Collection Permit (1823 to Caron). Funding was provided by the Royal Ontario
Museum Reproductions Acquisitions and Research Fund, Natural Sciences and
Engineering Research Council (NSERC) Discovery grants (311727 to Mángano,
341944 to Caron), and a grant from the Pomona College Faculty Research Fund.
Peter Fenton assisted with field work and Jason Loxton provided support in the
field. This is Royal Ontario Museum Burgess Shale Research Project number 26.
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Manuscript received 31 January 2010
Revised manuscript received 9 April 2010
Manuscript accepted 14 April 2010
Printed in USA
GEOLOGY, September 2010