The Burgess Shale

Phylas: Mollusca

Wiwaxia corrugata

3D animation of Wiwaxia corrugata grazing on Morania confluens.

ANIMATION BY PHLESCH BUBBLE © ROYAL ONTARIO MUSEUM

Taxonomy:

Kingdom: Mollusca
Phylum: Mollusca
Class: Unranked clade halwaxiids (stem group molluscs)
Species name: Wiwaxia corrugata
Remarks:

The relationship of Wiwaxia is hotly debated; its similarities to the molluscs have been highlighted (Conway Morris, 1985; Scheltema et al., 2003; Caron et al., 2006; Caron et al., 2007), but Matthew’s original view that it was related to the annelid worms (Matthew, 1899) still finds some adherents (Butterfield, 1990; Conway Morris and Peel, 1995; Butterfield, 2006; 2008). It is also possible that Wiwaxia branched off before the molluscs and annelids diverged (Eibye-Jacobsen, 2004). Wiwaxia has recently been placed in a group called the halwaxiids, along with the halkieriids, Orthrozanclus, and Odontogriphus (Conway Morris and Caron, 2007).

Described by: Matthew
Description date: 1899
Etymology:

Wiwaxia – from Wiwaxy Peaks (2,703 m) in Yoho National Park. The word wiwaxy is originally from the Stoney First Nation Nakoda language, meaning “windy.”

corrugata – from the Latin corrugis, “folded, or wrinkled,” in reference to the wrinkled aspect of the sclerites.

Type Specimens: Holotype –ROM8596 in the Royal Ontario Museum, Toronto, Canada.
Other species:

Burgess Shale and vicinity: none.

Other deposits: none described, although sclerites have been reported from a number of Middle Cambrian deposits extending from northern Canada (Butterfield, 1994) to China (Zhao et al., 1994).

Age & Localities:

Period:
Middle Cambrian, Bathyuriscus-Elrathina Zone (approximately 505 million years ago).
Principal localities:

The Walcott and Raymond Quarries on Fossil Ridge. The Trilobite Beds, Tulip Beds (S7) and Collins Quarry on Mount Stephen. Additional smaller localities are known on Mount Field and Mount Odaray.

History of Research:

Brief history of research:

In an early review of fossils collected from the Trilobite Beds on Mount Stephen by Walker, Canadian palaeontologist G. F. Matthew (1899) described several forms he thought represented tubes of various annelid worms, including one he named Orthotheca corrugata. At the time, Matthew did not know this particular fossil was only part of a much larger organism. It was only when Walcott (1911) discovered articulated and much better preserved specimens from the Phyllopod Bed that the morphology of this species became clearer. Walcott placed corrugata in his new genus Wiwaxia and interpreted it as a polychaete annelid worm (Walcott, 1911). The single best specimen of Walker’s “Orthotheca corrugata” remained unrecognized until it was “rediscovered” in the ROM collections in 1977.

Walcott’s interpretation was called into question in a comprehensive reassessment of the genus (Conway Morris, 1985), and Conway Morris’s link between Wiwaxia mouthparts and the molluscan radula was built upon by Scheltema et al. (2003) and Caron et al. (2006). Butterfield (1990), however, defended an annelid affinity mostly based on the study of individual sclerites, first at the crown-, and later at the stem-group level (Butterfield, 2003; 2006), but further work suggested that the evidence does not conclusively support a close relationship with annelids (Eibye-Jacobsen, 2004). A connection with the halkieriids was drawn early on (Bengtson and Morris, 1984; Conway Morris and Peel, 1995), and expanded more recently (Conway Morris and Caron, 2007).

Other studies have dealt more specifically with the ecology and taphonomy of this animal. The finely spaced patterning of ridges on the scale may have given Wiwaxia an iridescent aspect in life (Parker, 1998). Wiwaxia has proven useful in calculating the extent of decay in fossil assemblages (Caron and Jackson, 2006) and in reconstructing the longer term taphonomic processes responsible for the preservation of the Burgess Shale fossils (Butterfield et al., 2007).

Description:

Morphology:

Wiwaxia corrugata is a slug-like organism up to 5.5 cm in length almost entirely covered (except on the ventral surface) with an array of scale-like elements referred to as sclerites and spines. The body is roughly oval, and lacks evidence of segmentation. The body-covering sclerites are arranged in about 50 rows. In addition, two rows of 7–11 blade-like spines are present on the dorsal surface. Spines and sclerites were inserted directly into the body wall. Wiwaxia’s feeding apparatus consists of two (in rare cases three) toothed plates that have been compared to a molluscan radula or annelid jaws.

Abundance:

Wiwaxia is mostly known from the Walcott Quarry where it is relatively common, representing 0.9% of the specimens counted in the community (Caron and Jackson, 2008).

Maximum Size:
55 mm

Ecology:

Life habits: Mollusca
Feeding strategies: Mollusca
Ecological Interpretations:

The similarity of Wiwaxia’s feeding apparatus to that of Odontogriphus suggests that it too fed on the cyanobacterial Morania mats growing on the Cambrian sea floor. Its sclerite armour-plating and long spines, sometimes found broken, suggest that it was targeted by unidentified predators.

References:

BENGSTON, S. AND S. CONWAY MORRIS, 1984. A comparative study of Lower Cambrian Halkieria and Middle Cambrian Wiwaxia. Lethaia, 17:307-329.

BUTTERFIELD, N. J. 1990. A reassessment of the enigmatic Burgess Shale fossil Wiwaxia corrugata (Matthew) and its relationship to the polychaete Canadia spinosa Walcott. Paleobiology: 287-303.

BUTTERFIELD, N. J. 1994. Burgess Shale-type fossils from a Lower Cambrian shallow-shelf sequence in northwestern Canada. Nature, 369(6480): 477-479.

BUTTERFIELD, N. J. 2003. Exceptional fossil preservation and the Cambrian Explosion. Integrative and Comparative Biology, 43:166-177.

BUTTERFIELD, N. J. 2006. Hooking some stem-group “worms”: fossil lophotrochozoans in the Burgess Shale. BioEssays, 28: 1161-1166.

BUTTERFIELD, N. J. 2008. An early Cambrian radula. Journal of Paleontology, 82(3): 543-554.

CARON, J.-B. AND D. A. JACKSON, 2008. Paleoecology of the Greater Phyllopod Bed community, Burgess Shale. Palaeogeography, Palaeoclimatology, Palaeoecology, 258: 222-256.

CARON, J.-B., A. H. SCHELTEMA, C. SCHANDER AND D. RUDKIN, 2006. A soft-bodied mollusc with radula from the Middle Cambrian Burgess Shale. Nature, 442(7099): 159-163.

CARON, J.-B., A. H. SCHELTEMA, C. SCHANDER AND D. RUDKIN. 2007. Reply to Butterfield on stem-group “worms:” fossil lophotrochozoans in the Burgess Shale. BioEssays, 29:200-202.

CONWAY MORRIS, S. 1985. The Middle Cambrian metazoan Wiwaxia corrugata (Matthew) from the Burgess Shale and Ogygopsis Shale Shale, British Columbia, Canada. Philosophical Transactions of the Royal Society of London, Series B, 307(1134): 507-582.

CONWAY MORRIS, S. AND J.-B. CARON, 2007. Halwaxiids and the Early Evolution of the Lophotrochozoans. Science, 315(5816): 1255-1258.

CONWAY MORRIS, S. AND J. S. PEEL, 1995. Articulated halkieriids from the Lower Cambrian of North Greenland and their role in early protostome evolution. Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences, 347(1321): 305-358.

EIBYE-JACOBSEN, D. 2004. A reevaluation of Wiwaxia and the polychaetes of the Burgess Shale. Lethaia, 37(3): 317-335.

MATTHEW, G. F. 1899. Studies on Cambrian Faunas, No. 3. Upper Cambrian fauna, Mount Stephen, British Columbia. The trilobites and worms. Transactions of the Royal Society, 5: 39-66.

PARKER, A. R. 1998. Colour in Burgess Shale animals and the effect of light on evolution in the Cambrian. Proceedings of the Royal Society B: Biological Sciences, 265(1400): 967.

SCHELTEMA, A. H., K. KERTH AND A. M. KUZIRIAN, 2003. Original molluscan radula: Comparisons among Aplacophora, Polyplacophora, Gastropoda, and the Cambrian fossil Wiwaxia corrugata. Journal of Morphology, 257(2): 219-245.

WALCOTT, C. D. 1911. Middle Cambrian annelids. Smithsonian Miscellaneous Collections, 57(2): 109-144.

ZHAO, Y.-l., Y. QIAN AND X.-S. LI, 1994. Wiwaxia from Early-Middle Cambrian Kaili Formation in Taijiang, Guizhou. Acta Palaeontologica Sinica, 33:359-366.

Other Links:

http://www.paleobiology.si.edu/burgess/wiwaxia.html

Scenella amii

3D animation of Scenella amii.

ANIMATION BY PHLESCH BUBBLE © ROYAL ONTARIO MUSEUM

Taxonomy:

Kingdom: Mollusca
Phylum: Mollusca
Class: Unranked clade (stem group molluscs)
Species name: Scenella amii
Remarks:

Scenella is generally classified as a monoplacophoran mollusc (Knight, 1952; Runnegar and Jell, 1976). A position possibly ancestral to brachiopods (Dzik, 2010), or within the Cnidaria, has also been proposed (Babcock and Robison, 1988; Yochelson and Gil Cid, 1984).

Described by: Matthew
Description date: 1902
Etymology:

Scenella – from the Greek word skene, “tent, or shelter,” in reference to its shape.

amii – after Marc Henri Ami from the Geological Survey of Canada.

Type Specimens: Holotype –ROM8048 in the Royal Ontario Museum, Toronto, ON, Canada.
Other species:

Burgess Shale and vicinity: none

Other deposits: Dozens of species are known from the Lower Cambrian to the Lower Ordovician.

Age & Localities:

Period:
Middle Cambrian, Bathyuriscus-Elrathina Zone (approximately 505 million years ago).
Principal localities:

The Walcott and Raymond Quarries on Fossil Ridge. The Trilobite Beds and smaller localities on Mount Stephen.

History of Research:

Brief history of research:

The limpet-like appearance of Scenella led to its original classification as a mollusc, initially as a pteropod, then as a gastropod (Walcott, 1886). The first fossils of this genus known from the Burgess Shale were collected from the Trilobite Beds on Mount Stephen. These were described as Metoptoma amii by Matthew (1902), but Walcott (1908) considered other specimens from the same locality (and from the Walcott Quarry) to belong to Scenella varians, an earlier named species. Resser (1938) recognized that both species were identical and proposed a new combination, Scenella amii. In the same publication, Resser named a second species from the Trilobite Beds S. columbiana; this was based on a single specimen, originally recognized as a brachiopod with possible spines (Walcott, 1912), and remains highly dubious.

Description:

Morphology:

Each cone-shaped fossil has the form of a flat disc with a central peak, here termed “shell.” Concentric rings surround this peak, and sometimes the shell is also corrugated. The shells are stretched along one axis, making them elliptical rather than circular.

The fossils are often preserved in dense clusters and are usually oriented point-up.

No soft tissue is ever found associated with Scenella. The shell was evidently mineralized as indicated by the three-dimensional preservation and the presence of small cracks suggesting brittleness.

Abundance:

Hundreds of specimens of S. amii are known in the Walcott Quarry (2.27% of the community, Caron and Jackson, 2008). Many of these are found in dense clusters on single slabs.

Maximum Size:
10 mm

Ecology:

Life habits: Mollusca
Feeding strategies: Mollusca
Ecological Interpretations:

If a mollusc, Scenella would have been a creeping bottom-dweller, potentially a grazer.

References:

BABCOCK, L. E. AND R. A. ROBISON. 1988. Taxonomy and paleobiology of some Middle Cambrian Scenella (Cnidaria) and hyolithids (Mollusca) from western North America. University of Kansas Paleontological Contributions, Paper, 121: 1-22.

CARON, J.-B. AND D. A. JACKSON. 2008. Paleoecology of the Greater Phyllopod Bed community, Burgess Shale. Palaeogeography, Palaeoclimatology, Palaeoecology, 258: 222-256.

DZIK, J. 2010. Brachiopod identity of the alleged monoplacophoran ancestors of cephalopods. Malacologia, 52:97-113.

KNIGHT, J. B. 1952. Primitive fossil gastropods and their bearing on gastropod evolution. Smithsonian Miscellaneous Collections, 117(13): 1–56.

MATTHEW, G. F. 1902. Notes on Cambrian Faunas: Cambrian Brachiopoda and Mollusca of Mt. Stephen, B.C. with the description of a new species of Metoptoma. Transactions of the Royal Society of Canada, 4:107-112.

RASETTI, F. 1954. Internal shell structures in the Middle Cambrian gastropod Scenella and the problematic genus Stenothecoides. Journal of Paleontology, 28: 59-66.

RESSER, C. E. 1938. Fourth contribution to nomenclature of Cambrian fossils. Smithsonian Miscellaneous Collections, 97:1-43.

Runnegar, B. AND P. A. JELL. 1976. Australian Middle Cambrian molluscs and their bearing on early molluscan evolution. Alcheringa: An Australasian Journal of Palaeontology, 1(2): 109-138.

WALCOTT, C. D. 1886. Second contribution to the studies on the Cambrian faunas of North America. Bulletin of the United States Geological Survey, (30): 11-356.

WALCOTT, C. 1908. Mount Stephen rocks and fossils. Canadian Alpine Journal, 1: 232-248.

WALCOTT, C. 1912. Cambrian Brachiopoda. United States Geological Survey Monograph, 51: Part 1: 1-872, Part 872: 871-363.

YOCHELSON, E. L. AND D. GIL CID. 1984. Reevaluation of the systematic position of Scenella. Lethaia, 17: 331-340.

Other Links:

None

Haplophrentis carinatus

3D animation of Haplophrentis carinatus.

Animation by Phlesch Bubble © Royal Ontario Museum

Taxonomy:

Kingdom: Mollusca
Phylum: Mollusca
Class: Hyolitha (Order: Hyolithida, stem group molluscs)
Species name: Haplophrentis carinatus
Remarks:

Haplophrentis belongs to a group of enigmatic cone-shaped to tubular fossils called hyoliths that are known only from the Palaeozoic. Their taxonomic position is uncertain, but the Hyolitha have been regarded as a separate phylum, an extinct Class within Mollusca (Malinky and Yochelson, 2007), or as stem-group molluscs.

Described by: Matthew
Description date: 1899
Etymology:

Haplophrentis – from the Greek haploos, “single,” and phrentikos, “wall,” in reference to the single wall within the shell.

carinatus – from the Latin carinatus, “keel-shaped,” referring to the morphological similarity to the bottom of a boat.

Type Specimens: Lectotype –ROM8463a in the Royal Ontario Museum, Toronto, Canada.
Other species:

Burgess Shale and vicinity: none

Other deposits: H. reesei Babcock & Robinson, 1988 (type species), from the lower Middle Cambrian Spence Shale and elsewhere in Utah; H.? cf. carinatus from the Middle Cambrian Kaili deposit in China (Chen et al., 2003).

Age & Localities:

Period:
Middle Cambrian, Albertella Zone to Bathyuriscus-Elrathina Zone (approximately 505 million years ago).
Principal localities:

The Walcott, Raymond and Collins Quarries on Fossil Ridge, the Trilobite Beds on Mount Stephen and Stanley Glacier in Kootenay National Park.

History of Research:

Brief history of research:

Matthew described Hyolithes carinatus from the Trilobite Beds in 1899 based on five incomplete specimens. Babcock and Robison (1988) reviewed the original fossils, along with additional specimens collected by the Royal Ontario Museum from various Burgess Shale localities. They concluded that the species carinatus didn’t belong in Hyolithes, and established a new genus, Haplophrentis, to accommodate it.

Description:

Morphology:

Like all hyoliths, Haplophrentis had a weakly-mineralized skeleton that grew by accretion, consisting of a conical living shell (conch), capped with a clam-like “lid” (operculum), with two slender, curved and rigid structures known as “helens” protruding from the shell’s opening. The function of these helens is still debated, but one possibility was to allow settlement and stabilization on the sea floor. Haplophrentis had a wiggly gut whose preserved contents are similar to the surrounding mud.

H. carinatus usually grew to around 25 mm in length, although some specimens reached as much as 40 mm; the species is distinguished from H. reesei, its cousin from Utah, by the faint grooves on its upper surface, the more pronounced net-like pattern on its “lid” (operculum), and its wider, more broadly-angled living shell (conch).

Haplophrentis can be distinguished from the similar hyolith genus Hyolithes because it bears a longitudinal wall running down the inner surface of the top of its living-shell.

Abundance:

Haplophrentis is relatively common on Fossil Ridge and in the Walcott Quarry in particular, accounting for 0.35% of the community there (Caron and Jackson, 2008).

Maximum Size:
40 mm

Ecology:

Life habits: Mollusca
Feeding strategies: Mollusca
Ecological Interpretations:

Haplophrentis probably moved very little; its helens appear unsuited for use in locomotion (See Butterfield and Nicholas, 1996; Martí Mus and Bergström, 2005; Runnegar et al., 1975). Whilst Haplophrentis feeding mode remains somewhat conjectural, it probably consumed small organic particles from the seafloor. Numerous specimens have been found in aggregates or in the gut of the priapulid worm Ottoia prolifica suggesting Haplophrentis was actively preyed upon and ingested (Conway Morris, 1977; Babcock and Robison, 1988).

References:

BABCOCK, L. E. AND R. A. ROBISON. 1988. Taxonomy and paleobiology of some Middle Cambrian Scenella (Cnidaria) and hyolithids (Mollusca) from western North America. University of Kansas Paleontological Contributions, Paper, 121: 1-22.

BUTTERFIELD, N. J. AND C. NICHOLAS. 1996. Burgess Shale-type preservation of both non-mineralizing and “shelly” Cambrian organisms from the Mackenzie Mountains, Northwestern Canada. Journal of Paleontology, 70: 893-899.

CARON, J.-B. AND D. A. JACKSON. 2008. Paleoecology of the Greater Phyllopod Bed community, Burgess Shale. Palaeogeography, Palaeoclimatology, Palaeoecology, 258: 222-256.

CHEN, X. Y. ZHAO AND P. WANG. 2003. Preliminary research on hyolithids from the Kaili Biota, Guizhou. Acta Micropalaeontologica Sinica, 20: 296-302.

CONWAY MORRIS, S. 1977. Fossil priapulid worms. Special Papers in Palaeontology, 20: 1-95.

MALINKY, J. M. AND E. L. YOCHELSON. 2007. On the systematic position of the Hyolitha (Kingdom Animalia). Memoir of the Association of Australasian Palaeontologists, 34: 521-536.

MARTÍ MUS, M. AND J. BERGSTRÖM. 2005. The morphology of hyolithids and its functional implications. Palaeontology, 48:1139-1167.

MATTHEW, G. F. 1899. Studies on Cambrian faunas, No. 3. Upper Cambrian fauna of Mount Stephen, British Columbia. The trilobites and worms. Transactions of the Royal Society of Canada, Series 2, 4: 39-66.

RUNNEGAR, B., J. POJETA, N. J. MORRIS, J. D. TAYLOR, M. E. TAYLOR AND G. MCCLUNG. 1975. Biology of the Hyolitha. Lethaia, 8: 181-191.

Other Links:

http://paleobiology.si.edu/burgess/haplophrentis.html

http://www.kumip.ku.edu/cambrianlife/Utah-Mollusks-Hyolithids.html

Orthrozanclus reburrus

Taxonomy:

3D animation of Orthrozanclus reburrus.

ANIMATION BY PHLESCH BUBBLE © ROYAL ONTARIO MUSEUM

Kingdom: Mollusca
Phylum: Mollusca
Class: Unranked clade halwaxiids (stem group molluscs)
Species name: Orthrozanclus reburrus
Remarks:

The classification of this animal remains ambiguous but is thought to belong to the halwaxiids, a group including Wiwaxia and halkierids (Conway Morris and Caron, 2007). Two alternative positions were proposed for the halwaxiids, as either stem-group lophotrochozoans (a group which includes molluscs, annelids, and brachiopods) or as stem-group molluscs (see also Sigwart and Sutton, 2007).

Described by: Conway Morris and Caron
Description date: 2007
Etymology:

Orthrozanclus – from the Greek orthros, “dawn,” referring to its ancestral position, and zanclon, “sickle-like,” in reference to its long sclerites.

reburrus – from the Latin reburrus, “with bristling hair,” in reference to its hairy appearance.

Type Specimens: Holotype –ROM57197 in the Royal Ontario Museum, Toronto, Canada.
Other species:

Burgess Shale and vicinity: none.

Other deposits: none.

Age & Localities:

Period:
Middle Cambrian, Bathyuriscus-Elrathina Zone (approximately 505 million years ago).
Principal localities:

The Walcott Quarry on Fossil Ridge.

History of Research:

Brief history of research:

Two specimens were collected by Walcott but were never described. Between 1994 and 2000, the Royal Ontario Museum collected nine additional specimens from the Walcott Quarry. These were recognized as an unknown sclerite-bearing animal (Type C, Caron and Jackson 2006) allowing for a formal description of this new species (Conway Morris and Caron, 2007).

Description:

Morphology:

Specimens vary from 6 mm to 11.3 mm in length. The ventral side is flat, whereas the dorsal surface is rounded in cross-section, bearing three zones of sclerites and one anterior shell. A set of relatively small sclerites is present around the margins of the body. These appear flat and slightly curved in one direction. Above this marginal set is a second set of much longer sclerites that seem to originate from a narrow zone along the entire length of the body. These are circular in cross section, appear to have a larger base, and tend to be curved and pointing upwards; they are probably hollow and bear one or two ridges. The presence of kinked sclerites suggests a lack of mineralization. A third set of much smaller sclerites covers the convex dorsal side of the body, but these are not clearly preserved. The shell, which was presumably mineralized, is triangular in outline with the pointed end towards the front. Fine striations along the shell are probably growth lines, indicating that growth occurred from the front to the back.

Abundance:

Very rare; all 11 known specimens come from the Walcott Quarry only, where it represents a tiny fraction of the community (0.02%) (Caron and Jackson, 2008).

Maximum Size:
11 mm

Ecology:

Life habits: Mollusca
Feeding strategies: Mollusca
Ecological Interpretations:

Orthrozanclus is similar in overall aspect and probably in ecology to the better known Wiwaxia corrugata.Orthrozanclus, like Wiwaxia, was probably herbivorous and would have crept along the seafloor in search for food.

References:

CARON, J.-B. AND D. A. JACKSON. 2008. Paleoecology of the Greater Phyllopod Bed community, Burgess Shale. Palaeogeography, Palaeoclimatology, Palaeoecology, 258: 222-256.

CONWAY MORRIS, S. AND J.-B. CARON. 2007. Halwaxiids and the early evolution of the lophotrochozoans. Science, 315: 1255-1258.

SIGWART, J. D. AND M. D. SUTTON. 2007. Deep molluscan phylogeny: synthesis of palaeontological and neontological data. Proceedings of the Royal Society B: Biological Sciences, 274: 2413-2419.

Other Links:

http://www.sciencemag.org/content/315/5816/1255

Oikozetetes seilacheri

Oikozetetes seilacheri (USNM 276247a) – Holotype (left); (GSC 110395) – Paratype (right). Morphs A and B respectively, represented at opposite ends of a presumed slug-like animal. Morph A length = 2.5 mm, Morph B length = 3.5 mm. Specimen dry – direct light. Walcott Quarry.

© SMITHSONIAN INSTITUTION – NATIONAL MUSEUM OF NATURAL HISTORY (LEFT) AND GEOLOGICAL SURVEY OF CANADA (RIGHT). PHOTOS: JEAN-BERNARD CARON

Taxonomy:

Kingdom: Mollusca
Phylum: Mollusca
Class: Unranked clade halwaxiids (stem group molluscs)
Species name: Oikozetetes seilacheri
Remarks:

Oikozetetes is thought to represent a halkieriid (Conway Morris, 1995). These organisms are generally considered to be related to the molluscs, although they may fall deeper in the lophotrochozoan stem (Conway Morris and Caron, 2007).

Described by: Conway Morris
Description date: 1995
Etymology:

Oikozetetes – from the Greek oikos, “home,” and the Latin zetetes, “quest,” referring to the challenge of classifying the fossils in their correct taxonomic “home.”

seilacheri – after the German paleontologist Adolf Seilacher.

Type Specimens: Holotype – UNSM 276247a in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.
Other species:

Burgess Shale and vicinity: none

Other deposits: One unnamed species has been reported from the Lower Cambrian Mernmerna Formation, Flinders Ranges, South Australia (Paterson et al., 2009).

Age & Localities:

Period:
Middle Cambrian, Bathyuriscus-Elrathina Zone (approximately 505 million years ago).
Principal localities:

The Walcott Quarry on Fossil Ridge.

History of Research:

Brief history of research:

No research has been performed on the Burgess Shale Oikozetetes since its initial description by Conway Morris in 1995. Conway Morris noted the similarity of the shells to those of the recently-discovered Halkieria (Conway Morris and Peel, 1990, 1995), and proposed that the two distinct shell forms both belonged to a single animal. Since then, three Oikozetetes-like shells have been found alongside mineralized Halkieria-type sclerites in Australian sediments (Paterson et al., 2009).

Description:

Morphology:

All that is known of Oikozetetes is its two mineralized shells (morphs A and B), which typically grow to about 6 mm. The shells are thought to have sat at either end of a slug-like animal similar to Halkieria. This organism may have been covered in Wiwaxia-like sclerites, or may have simply had a tough skin that is never preserved. The frontal shell (morph A) resembles a scallop, with concentric ridges probably representing growth lines surrounding its posterior point; larger flanges form an arrowhead pointing towards the rear of the organism. The rear shell (morph B) is steeply arched and D-shaped, again bearing concentric ridges.

Abundance:

Oikozetetes is rare and is often misidentified in the collections as fragments of the animal Scenella. Conway Morris’s (1995) description included 29 specimens.

Maximum Size:
10 mm

Ecology:

Life habits: Mollusca
Feeding strategies: Mollusca
Ecological Interpretations:

As with other Burgess Shale slug-like forms, Oikozetetes probably fed on the cyanobacterial Morania mats that grew on the Cambrian sea floor (Caron et al., 2006).

References:

CARON, J.-B., A. H. SCHELTEMA, C. SCHANDER AND D. RUDKIN. 2006. A soft-bodied mollusc with radula from the Middle Cambrian Burgess Shale. Nature, 442: 159-163.

CONWAY MORRIS, S. 1995. Enigmatic shells, possibly halkieriid, from the Middle Cambrian Burgess Shale, British Columbia. Neues Jahrbuch für Geologie und Palaeontologie. Abhandlungen, 195: 319-331.

CONWAY MORRIS, S. AND J. S. PEEL. 1990. Articulated halkieriids from the Lower Cambrian of north Greenland. Nature, 345: 802-805.

CONWAY MORRIS, S. AND J. S. PEEL. 1995. Articulated halkieriids from the Lower Cambrian of North Greenland and their role in early protostome evolution. Philosophical Transactions of the Royal Society of London Series B, 347: 305-358.

PATERSON, J. R., G. A. BROCK AND C. B. SKOVSTED. 2009. Oikozetetes from the early Cambrian of South Australia: implications for halkieriid affinities and functional morphology. Lethaia, 42: 199-203.

Other Links:

None

Odontogriphus omalus

3D animation of Odontogriphus omalus.

ANIMATION BY PHLESCH BUBBLE © ROYAL ONTARIO MUSEUM

Taxonomy:

Kingdom: Mollusca
Phylum: Mollusca
Class: Unranked clade halwaxiids (stem group molluscs)
Species name: Odontogriphus omalus
Remarks:

Odontogriphus is an early stem-group mollusc (Caron et al., 2006; Sigwart and Sutton, 2007), or a stem-group to the lophotrochozoans, a group which includes molluscs, annelids, and brachiopods (Conway Morris and Caron, 2007). A relationship to annelids (Butterfield, 2006) has been suggested, but appears less likely (Caron et al., 2007).

Described by: Conway Morris
Description date: 1976
Etymology:

Odontogriphus – from the Greek odontos, “tooth,” and griphos, “puzzle, or riddle,” in reference to its uncertain affinities.

omalus – from the Greek homalos, “flat,” in reference to the animal’s flattened shape.

Type Specimens: Holotype –USNM196169 in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.
Other species:

Burgess Shale and vicinity: none.

Other deposits: none.

Age & Localities:

Period:
Middle Cambrian, Bathyuriscus-Elrathina Zone (approximately 505 million years ago).
Principal localities:

The Walcott Quarry on Fossil Ridge and the Tulip Beds (S7) on Mount Stephen.

History of Research:

Brief history of research:

Walcott collected the first specimen between 1909 and 1924 but it remained unstudied for more than half a century. Conway Morris “rediscovered” the part and counterpart of this specimen in different sections of the Walcott collection and described it in 1976 as Odontogriphus omalus. The affinities of Odontogriphus remained uncertain until the Royal Ontario Museum discovered 189 new specimens between 1990 and 2000, allowing for a thorough redescription of the animal (Caron et al., 2006).

Description:

Morphology:

This entirely soft-bodied animal is ovoid and dorsoventrally compressed, reaching up to 125 mm in length and 43 mm in width. The front and back are semicircular in outline and of similar size. The mouth is ventral with a radula composed of two primary tooth rows. A muscular foot extends from behind the mouth to the posterior part of the animal and is surrounded by gills (or ctenidia), except at the front. The dorsal surface is smooth and does not bear any shells, spines or plates. Internally, a large stomach is preserved with a narrow and straight intestine ending in a sub-terminal anus.

Abundance:

Most specimens come from the Walcott Quarry, where Odontogriphus represents 0.42% of the community (Caron and Jackson, 2008). A single specimen comes from Mount Stephen (S7 locality).

Maximum Size:
125 mm

Ecology:

Life habits: Mollusca
Feeding strategies: Mollusca
Ecological Interpretations:

The presence of a radula suggests that Odontogriphus was a grazer, using its teeth to rasp and ingest food. Locomotory waves within the large foot would have enabled the animal to crawl along the surface of the mud. Odontogriphus might have fed on benthic, sheet-like masses of the cyanobacterium Morania, since fossils of both are often found associated in the same layers.

References:

BUTTERFIELD, N. J. 2006. Hooking some stem-group “worms”: fossil lophotrochozoans in the Burgess Shale. BioEssays, 28: 1161-1166.

CARON, J.-B., A. H. SCHELTEMA, C. SCHANDER AND D. RUDKIN. 2006. A soft-bodied mollusc with radula from the Middle Cambrian Burgess Shale. Nature, 442: 159-163.

CARON, J.-B., A. H. SCHELTEMA, C. SCHANDER AND D. RUDKIN. 2007. Reply to Butterfield on stem-group “worms:” fossil lophotrochozoans in the Burgess Shale. BioEssays, 29: 200-202.

CONWAY MORRIS, S. 1976. A new Cambrian lophophorate from the Burgess Shale of British Columbia. Palaeontology, 19: 199-222.

SIGWART, J. D. AND M. D. SUTTON. 2007. Deep molluscan phylogeny: synthesis of palaeontological and neontological data. Proceedings of the Royal Society B: Biological Sciences, 274: 2413-2419.

Other Links:

http://www.nature.com/nature/journal/v442/n7099/full/nature04894.html

Nectocaris pteryx

3D animation of Nectocaris pteryx.

ANIMATION BY PHLESCH BUBBLE © ROYAL ONTARIO MUSEUM

Taxonomy:

Kingdom: Mollusca
Phylum: Mollusca
Class: Cephalopoda (stem group molluscs)
Species name: Nectocaris pteryx
Remarks:

Nectocaris is regarded as an early stem-group mollusc close to the cephalopods. This stem-group also includes Vetustovermis from the Middle Cambrian Emu Bay Shale of Australia, and the Lower Cambrian Petalilium from the Chengjiang deposit in China (Smith and Caron, 2010).

Described by: Conway Morris
Description date: 1976
Etymology:

Nectocaris – from the Greek nekto, “swimming,” and the Latin caris, “shrimp,” based on its original interpretation as an arthropod.

pteryx – from the Greek pteryx, “fins,” in reference to the presence of fins.

Type Specimens: Holotype –USNM198667 in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.
Other species:

Burgess Shale and vicinity: none.

Other deposits: none.

Age & Localities:

Period:
Middle Cambrian, Bathyuriscus-Elrathina Zone (approximately 505 million years ago).
Principal localities:

The Walcott, Raymond and Collins Quarries on Fossil Ridge.

History of Research:

Brief history of research:

As with Odontogriphus, another Burgess Shale animal related to molluscs, Walcott collected the first specimen of Nectocaris between 1909 and 1924. The fossil was photographed by Walcott, and its print sat with the unidentified specimen in the Smithsonian collections until noticed and described by Simon Conway Morris in 1976. Due to the lateral compression of the fossil, his resulting reconstruction was laterally-oriented. The funnel, bent back over the front, resembled the head-shield of an arthropod, and yet the fin, folded along the top of the organism, looked much like the ray-bearing dorsal fin of a chordate. A chordate affinity was further suggested by the myomere-like appearance of the bars, and although Conway Morris did not offer a firm diagnosis, Simonetta (1988) promoted a chordate status (Insom et al., 1995).

Meanwhile, Glaessner had described Vetustovermis, based on an ill-preserved specimen from Australia’s Emu Bay Shale, and because of its segmented appearance he suggested an affinity with annelid worms (Glaessner, 1979). Other workers noted the similarity of some Chengjiang fossils to this specimen and described them as slug-like relatives of the molluscs (Chen et al., 2005). During this period, the Royal Ontario Museum had been collecting similar fossils, which Desmond Collins recognized as representatives of Nectocaris. These were eventually described as stem-group cephalopods (Smith and Caron, 2010). The relationships among members of this clade are difficult to determine, and it may require further fossil finds to establish their diversity and range. The absence of a shell in Nectocarisindicates that cephalopods, which were previously thought to have evolved later in the Cambrian from snail-like monoplacophorans, did not require a buoyant shell to start swimming, but derived their shell independently of other mollusc lineages.

Description:

Morphology:

The body of Nectocaris is kite-shaped and can reach up to 72 mm in length, including two flexible tentacles that extend forwards from the head, which also bears a pair of camera-type eyes on short stalks. A long, nozzle-like funnel originates under the base of the head. The main body has wide lateral fins with transverse bars; a large axial cavity contains paired gills.

Abundance:

Nectocaris is known from 90 specimens on Fossil Ridge, mostly from the Collins Quarry; it is rare or absent at most other Burgess Shale localities. Only two specimens, including the holotype, have been found in the Walcott Quarry.

Maximum Size:
72 mm

Ecology:

Life habits: Mollusca
Feeding strategies: Mollusca
Ecological Interpretations:

A free-swimming predator or scavenger, Nectocaris would have fed on small prey items with its prehensile tentacles in a similar fashion to squid today. Its primary mode of propulsion would have been in the flexing of its fins; it may have supplemented this by squirting water from its funnel. The funnel was also used to inhale and exhale water, which entered the animal’s body cavity to oxygenate the large internal gills.

References:

CHEN, J.-Y., D.-Y. HUANG AND D. J. BOTTJER. 2005. An Early Cambrian problematic fossil: Vetustovermis and its possible affinities. Proceedings of the Royal Society B: Biological Sciences, 272(1576): 2003-2007.

CONWAY MORRIS, S. 1976. Nectocaris pteryx, a new organism from the Middle Cambrian Burgess Shale of British Columbia. Neues Jahrbuch für Geologie und Paläontologie, Monatshefte, 12: 703-713.

GLAESSNER, M. F. 1979. Lower Cambrian Crustacea and annelid worms from Kangaroo Island, South Australia. Alcheringa, 3(1): 21-31.

INSOM, E. A. PUCCI AND A. M. SIMONETTA. 1995. Cambrian Protochordata, their origin and significance. Bollettino di Zoologia, 62(3): 243-252.

SIMONETTA, A. M. 1988. Is Nectocaris pteryx a chordate? Bollettino di Zoologia, 55(1-2): 63-68.

SMITH, M. AND J.-B. CARON. 2010. Primitive soft-bodied cephalopods from the Cambrian. Nature, 465: 469-472.

Other Links:

http://www.nature.com/nature/journal/v465/n7297/full/nature09068.html