The Burgess Shale

Vauxia gracilenta

3D animation of Vauxia bellula and other sponges (Choia ridleyiDiagoniella cyathiformisEiffelia globosaHazelia confertaPirania muricata, and Wapkia elongata) and Chancelloria eros a sponge-like form covered of star-shaped spines.

ANIMATION BY PHLESCH BUBBLE © ROYAL ONTARIO MUSEUM

Taxonomy:

Class: Demospongea (Order: Verongida)
Remarks:

Vauxia was placed within the hexactinellids by Walcott in his 1920 original description but Rigby (1980) transferred the genus and family to the Demospongea. Demosponges, the same group that are harvested as bath sponges, represent the largest class of sponges today.

Species name: Vauxia gracilenta
Described by: Walcott
Description date: 1920
Etymology:

Vauxia – from Mount Vaux (3,319 m), a mountain Peak in Yoho National Park, British Columbia. The name refers to William Sandys Wright Vaux (1818-1885) an antiquarian at the British Museum.

gracilenta – from the Latin gracilis, “slender,” referring to the delicate structure of the sponge.

Type Specimens: Lectotypes –USNM66515 (V. gracilenta),USNM66508 (V. bellula),USNM66517 (V. densa),USNM66520 (V. venata), in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA. Holotype –ROM53572 (V. irregulara) in the Royal Ontario Museum, Toronto, Canada.
Other species:

Burgess Shale and vicinity: V. bellula Walcott, 1920; V. densa Walcott, 1920; V. irregulara Rigby and Collins, 2004; V. venata Walcott, 1920.

Other deposits: none.

Age & Localities:

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

Burgess Shale and vicinity: Vauxia species are known in the Walcott, Raymond and Collins Quarries on Fossil Ridge, the Trilobite Beds, Tulip Beds (S7) and the Collins Quarry on Mount Stephen, and smaller sites on Mount Field and Odaray Mountain. Vauxia is also known from Monarch in Kootenay National Park.

Other deposits: V. bellula Walcott, 1920 from the Middle Cambrian Wheeler and Marjum Formations in Utah (Rigby et al., 2010); V. magna Rigby, 1980 from the Middle Cambrian Spence Shale in Utah (Rigby, 1980).

History of Research:

Brief history of research:

This sponge was originally described by Walcott in 1920. The genus was reviewed by Rigby (1980) and the species redescribed by Rigby (1986) and Rigby and Collins (2004) in their examination of the Burgess Shale sponges.

Description:

Morphology:

Specimens of Vauxia gracilenta can range from simple unbranched forms to more complex branching forms and reach up to 8 cm in height. Each branch is deeply conical and almost cylindrical, with a simple open central cavity (spongocoel) ending in a rounded of flat opening (osculum). The skeleton is double layered with a thin dermal layer and an inner layer (endosomal). The dermal layer has small openings (ostia) and is composed of a dense network of ladder-like fibers supported by radial fibers from the inner layer. The inner layer forms a regular reticulated net-like skeleton of fibers with 4-6 sided polygons which is characteristic of the genus and species. The fibrous elements (spongin) represent tough collagen proteins. There is no evidence of siliceous spicules in the skeleton.

The different species have been identified mostly based on variations of the skeletal elements and the shape of the branches. Some species can reach up to at least 15 cm in height (V. bellulaV. densa).

Abundance:

Vauxia is relatively common in the Raymond Quarry and other sites on Mount Stephen but is rare in the Walcott Quarry where it represents less than 0.05% of the community (Caron and Jackson, 2008).

Maximum Size:
80 mm

Ecology:

Ecological Interpretations:

Vauxia would have lived attached to the sea floor. Particles of organic matter were extracted from the water as they passed through canals in the sponge’s wall.

References:

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

RIGBY, J. K. 1980. The new Middle Cambrian sponge Vauxia magna from the Spence Shale of Northern Utah and taxonomic position of the Vauxiidae. Journal of Paleontology, 54(1): 234-240.

RIGBY, J. K. 1986. Sponges of the Burgess Shale (Middle Cambrian), British Columbia. Palaeontographica Canadiana, 2: 1-105 p.

RIGBY, J. K. AND D. COLLINS. 2004. Sponges of the Middle Cambrian Burgess Shale and Stephen Formations, British Columbia. Royal Ontario Museum Contributions in Science (1): 155 p.

RIGBY, J. K., S. B. CHURCH AND N. K. ANDERSON. 2010. Middle Cambrian Sponges from the Drum Mountains and House Range in Western Utah. Journal of Paleontology, 84: 66-78.

WALCOTT, C. D. 1920. Middle Cambrian Spongiae. Cambrian Geology and Paleontology IV. Smithsonian Miscellaneous Collections, 67(6): 261-365.

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Pirania muricata

3D animation of Pirania muricata and other sponges (Choia ridleyi, Diagoniella cyathiformis, Eiffelia globosa, Hazelia conferta, Vauxia bellula, and Wapkia elongata) and Chancelloria eros a sponge-like form covered of star-shaped spines.

ANIMATION BY PHLESCH BUBBLE © ROYAL ONTARIO MUSEUM

Taxonomy:

Class: Demospongea (Order: Monaxonida)
Remarks:

Pirania is considered a primitive demosponge (Rigby, 1986). Demosponges, the same group that are harvested as bath sponges, represent the largest class of sponges today.

Species name: Pirania muricata
Described by: Walcott
Description date: 1920
Etymology:

Pirania – from Mount Saint Piran (2,649 m), situated in the Bow River Valley in Banff National Park, Alberta. Samuel Allen named Mount St. Piran after the Patron Saint of Cornwall in 1894.

muricata – from the Latin muricatus, “pointed, or full of sharp points.” The name refers to the large pointed spicules extending out from the wall of the sponge.

Type Specimens: Lectotype –USNM66495 (erroneously referred as 66496 in Rigby, 1986), in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.
Other species:

Burgess Shale and vicinity: none

Other deposits: Pirania auraeum Botting, 2007 from the Lower Ordovician of Morocco (Botting, 2007); Pirania llanfawrensis Botting, 2004 from the Upper Ordovician of England (Botting, 2004).

Age & Localities:

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

The Walcott Quarry on Fossil Ridge. The Trilobite Beds and Tulip Beds (S7) on Mount Stephen and several smaller sites on Mount Field, Mount Stephen and Mount Odaray.

History of Research:

Brief history of research:

Pirania was first described by Walcott (1920). Rigby (1986) redescribed this sponge and concluded that the skeleton is composed of hexagonally arranged canals, large pointed spicules and tufts of small spicules. This sponge was also reviewed by Rigby and Collins based on new material collected by the Royal Ontario Museum (2004).

Description:

Morphology:

Pirania is a thick-walled cylindrical sponge that can have up to four branches. The skeleton of the sponge is composed of tufts of small spicules and has very distinctive long pointed spicules that emerge from the external wall. Long canals perforate the wall of the sponge to allow water flow through it. Branching occurs close to the base of the sponge.

Abundance:

Pirania is common in most Burgess Shale sites but comprises only 0.38% of the Walcott Quarry community (Caron and Jackson, 2008).

Maximum Size:
30 mm

Ecology:

Ecological Interpretations:

Pirania would have lived attached to the sea floor. Particles of organic matter were extracted from the water as they passed through canals in the sponge’s wall. The brachiopods Nisusia and Micromitra a range of other sponges and even juvenile chancelloriids are often found attached to the long spicules of this sponge, possibly to avoid higher turbidity levels near the seafloor.

References:

BOTTING, J. P. 2004. An exceptional Caradoc sponge fauna from the Llanfawr Quarries, Central Wales and phylogenetic implications. Journal of Systematic Paleontology, 2: 31-63.

BOTTING, J. P. 2007. ‘Cambrian’ demosponges in the Ordovician of Morocco: insights into the early evolutionary history of sponges. Geobios, 40: 737-748.

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

RIGBY, J. K. 1986. Sponges of the Burgess shale (Middle Cambrian), British Columbia. Palaeontographica canadiana, 2: 105 p.

RIGBY, J. K. AND D. COLLINS. 2004. Sponges of the Middle Cambrian Burgess Shale and Stephen Formations, British Columbia. Royal Ontario Museum Contributions in Science (1): 155 p.

WALCOTT, C. D. 1920. Middle Cambrian Spongiae. Cambrian Geology and Paleontology IV. Smithsonian Miscellaneous Collections, 67(6): 261-365.

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Paterina zenobia

3D animation of Paterina zenobia and other brachiopods (Acrothyra gregaria, Diraphora bellicostata, Micromitra burgessensis, and Nisusia burgessensis).

ANIMATION BY PHLESCH BUBBLE © ROYAL ONTARIO MUSEUM

Taxonomy:

Class: Paterinata (Order: Paterinida)
Remarks:

A brachiopod within the family Paterinidae.

Species name: Paterina zenobia
Described by: Walcott
Description date: 1912
Etymology:

Paterina – from the Latin word pater, “father,” because the species was considered the ancestor of modern brachiopods, and the diminutive suffix, – ina, “derived from.”

zenobia – possibly from the Greek, Zeon, a form of Zeus.

Type Specimens: Syntype–USNM58311; plesiotypesUSNM56907, 51483, 69631- 69637 in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.
Other species:

Burgess Shale and vicinity: None to date. The Burgess Shale brachiopods, in particular from the Trilobite Beds on Mount Stephen, need to be re-examined (see also Brief history of research).

Other deposits: Several species are known in the Lower to the Middle Cambrian worldwide.

Age & Localities:

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

The Walcott Quarry on Fossil Ridge and the Trilobite Beds on Mount Stephen.

History of Research:

Brief history of research:

Walcott originally assigned specimens collected from the Burgess Shale and Mount Stephen to Micromitra zenobia Walcott (1912) and a subspecies of Paterina stissingensis, called Paterina stissingensis ora Walcott (1912). Both taxa were redescribed as Paterina zenobia by Resser (1938), a combination still in use today. However, close similarities between species of the two genera have created difficulties in defining their specific characteristics, which have resulted in many incorrectly identified specimens.

Description:

Morphology:

Paterina is the type genus of one of the earliest and most primitive brachiopod groups, the Paterinata. Unlike many modern brachiopods, its hinge line is straight and crosses almost the full width of the shell. The moderately biconvex shell grows consistently, rather than showing separate stages of development. Its exterior growth lines are coarse and regular. Faint radial ridges are present at the apex of some adult specimens. No preserved soft parts are known and the shell was originally mineralized.

Abundance:

This species is rare in the Walcott Quarry and represents a very small fraction of the entire fauna (<0.05%) (Caron and Jackson, 2008).

Maximum Size:
11 mm

Ecology:

Ecological Interpretations:

Paterina probably attached to the substrate by a very short stalk. Paterina extracted food particles from the water with its filter-feeding apparatus (located between the shells) called a lophophore.

References:

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

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

WALCOTT, C. D. 1912. Cambrian Brachiopoda. United States Geological Survey, Monograph, 51: part I, 812 p; part II, 363 p.

Other Links:

None

Nisusia burgessensis

3D animation of Nisusia burgessensis and other brachiopods (Acrothyra gregaria, Diraphora bellicostata, Micromitra burgessensis, and Paterina zenobia).

ANIMATION BY PHLESCH BUBBLE © ROYAL ONTARIO MUSEUM

Taxonomy:

Class: Kutorginata (Order: Kutorginida)
Remarks:

Nisusia belongs within the Family Nisusiidae.

Species name: Nisusia burgessensis
Described by: Walcott
Description date: 1889
Etymology:

Nisusia – from the Latin, nisus, meaning “labored, or striven.”

burgessensis – from Mount Burgess (2,599 m), a mountain peak in Yoho National Park. Mount Burgess was named in 1886 by Otto Klotz, the Dominion topographical surveyor, after Alexander Burgess, a former Deputy Minister of the Department of the Interior.

Type Specimens: Syntypes –USNM69690-69697 in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.
Other species:

Burgess Shale and vicinity: Nisusia alberta from the Trilobite Beds on Mount Stephen (Walcott, 1905, 1908). The Burgess Shale brachiopods, in particular from the Trilobite Beds on Mount Stephen, need to be re-examined (see also Brief history of research).

Other deposits: Several species are known in the Lower-Middle Cambrian of North America, Greenland, Russia, China and Australia.

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:

Nisusia burgessensis was originally described as Orthisina alberta (Walcott, 1889) before being renamed Nisusia alberta (Walcott, 1905). Specimens of this species identified from the Walcott Quarry (Walcott, 1912) were re-described by Walcott as Nisusia burgessensis(Walcott, 1924), a combination still in use today. This species has not been studied since and is in need of revision.

Description:

Morphology:

This species has fine radiating ornamental lines (costae) and concentric lines of growth. The shell was originally mineralized. It is roughly 1.5 wider than its length. Both valves are convex, but the convexity of the ventral shell is more pronounced. The shells would have been articulated with short and small teeth, like in Diraphora, a comparable form from the Burgess Shale. Very thin bristles (setae) are present in a single specimen at the front of the shell margin. These would have been attached to the edge of the mantle along both the dorsal and ventral valves in the same way as in Micromitra.

Abundance:

Nisusia burgessensis is relatively common in the Walcott Quarry but overall represents a small fraction of the fauna (<0.3%) (Caron and Jackson, 2008).

Maximum Size:
23 mm

Ecology:

Ecological Interpretations:

Nisusia probably had a relatively short, stout pedicle attached either to the substrate or to other organisms like the sponge Pirania, to raise it above the sediment-water interface. In this way the brachiopod would have been relatively protected from flocculent mud travelling along the sediment-water interface, which could have been detrimental to its filter-feeding apparatus (located between the shells) called a lophophore. The bristles (setae) might have also helped reduce the intake of mud particles into the filter-feeding apparatus.

References:

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

WALCOTT, C. 1889. Description of new genera and species of fossils from the Middle Cambrian. United States National Museum, Proceedings for 1888: 441-446.

WALCOTT, C. 1905. Cambrian brachiopods with descriptions of new genera and species. United States National Museum, Proceedings for 1905: 227-337.

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

WALCOTT, C. D. 1912. Cambrian Brachiopoda. United States Geological Survey, Monograph, 51: part I, 812 p; part II, 363 p.

WALCOTT, C. D. 1924. Cambrian and Ozarkian Brachiopoda. Cambrian Geology and Paleontology IV. Smithsonian Miscellaneous Publications, 67: 477-554.

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Micromitra burgessensis

3D animation of Micromitra burgessensis and other brachiopods (Acrothyra gregaria, Diraphora bellicostata, Nisusia burgessensis, and Paterina zenobia).

ANIMATION BY PHLESCH BUBBLE © ROYAL ONTARIO MUSEUM

Taxonomy:

Class: Paterinata (Order: Paterinida)
Remarks:

Micromitra belongs within the Family Paterinidae.

Species name: Micromitra burgessensis
Described by: Walcott
Description date: 1908
Etymology:

Micromitra – from the Greek mikros, “small,” and mitra, “turban.”

burgessensis – from Mount Burgess (2,599 m), a mountain peak in Yoho National Park. Mount Burgess was named in 1886 by Otto Klotz, the Dominion topographical surveyor, after Alexander Burgess, a former Deputy Minister of the Department of the Interior.

Type Specimens: Holotype –USNM69646 in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.
Other species:

Burgess Shale and vicinity: none to date. The Burgess Shale brachiopods, in particular from the Trilobite Beds on Mount Stephen, need to be re-examined (see also Brief history of research).

Other deposits: Numerous species, all from the Cambrian, are known worldwide.

Age & Localities:

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

The Walcott and Raymond Quarries on Fossil Ridge. Additional localities are known on Mount Field, Mount Stephen, and near Stanley Glacier.

History of Research:

Brief history of research:

Originally identified as Micromitra (Iphidella) pannula by Walcott (1908) from the Trilobite Beds on Mount Stephen (see also Walcott, 1912), it was redescribed as a new species by Resser (1938). Resser’s description fails to distinguish Micromitra burgessensis from any other species of the genus, it was based upon only a single valve, and it was not illustrated. The validity of this species is questionable and needs reassessment.

Description:

Morphology:

This species is the most ornamented of the Burgess Shale brachiopods. The shell was originally mineralized. It has pronounced growth lines and fine raised lines which cut obliquely across the shell. The intersection between the different lines creates small diamonds on the surface of the shell. The valves are subcircular with the hinge nearly straight. Perhaps the most striking of the preserved features of this animal are long and slender bristles (setae) which extend far beyond the margins of the shell. These would have been attached to the edge of the mantle along both the dorsal and ventral valves.

Abundance:

Micromitra burgessensis is relatively common in the Walcott Quarry but overall represents a small fraction of the fauna (<0.3%) (Caron and Jackson, 2008). This species is also present in the Raymond Quarry on Fossil Ridge.

Maximum Size:
10 mm

Ecology:

Ecological Interpretations:

Many specimens of Micromitra burgessensis are preserved attached to spicules of the sponge Pirania, suggesting that this species was epibenthic, supported above the sediment-water interface. In this way the brachiopod would have been relatively protected from flocculent mud travelling along the sediment-water interface, which could have been detrimental to its filter-feeding apparatus (located between the shells) called a lophophore – The bristles might have also helped reduce mud particles.

References:

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

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

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

WALCOTT, C. D. 1912. Cambrian Brachiopoda. United States Geological Survey, Monograph, 51: part I, 812 p; part II, 363 p.

Other Links:

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Mackenzia costalis

3D animation of Mackenzia costalis.

ANIMATION BY PHLESCH BUBBLE © ROYAL ONTARIO MUSEUM

Taxonomy:

Class: Anthozoa? (Order: Actiniaria(?), stem group cnidarians)
Remarks:

Mackenzia is thought to be a cnidarian (a group which includes modern coral and jellyfish) and appears most similar to modern sea anemones (Conway Morris, 1993).

Species name: Mackenzia costalis
Described by: Walcott
Description date: 1911
Etymology:

Mackenzia – from Mount Mackenzie (2,461 m) near Revelstoke, southwest of the Burgess Shale. Mount Mackenzie was named in honor of Alexander Mackenzie (1822-1892), Canada’s 2nd Prime Minister.

costalis – from the Latin costalis, “pertaining to ribs.” The name refers to the lineations along the length of the animal.

Type Specimens: Lectotype –USNM57556 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 and Raymond Quarries on Fossil Ridge. The Tulip Beds (S7) on Mount Stephen.

History of Research:

Brief history of research:

Mackenzia was first described as a holothurian, a group of echinoderms commonly known as the sea-cucumbers (Walcott, 1911). Additional fossils collected by the Geological Survey of Canada and restudy of Walcott’s collection led Conway Morris (1989, 1993) to reinterpret this animal as a cnidarian.

Description:

Morphology:

Mackenzia is a large saclike animal, up to 16 cm in height, which was anchored to hard substrates with a disc or holdfast via a short stalk; it probably stood upright. The surface of the body is folded longitudinally into 8-10 ridges. There is a large gut cavity and some evidence of internal partitioning, but little else is known of the anatomy. Tentacles are absent; the mouth was probably at the end opposite the stalk.

Abundance:

Mackenzia is very rare and represents only 0.03% of the Walcott Quarry community (Caron and Jackson, 2008).

Maximum Size:
200 mm

Ecology:

Ecological Interpretations:

Mackenzia probably lived on the seabed and may have attached to animal remains such as brachiopod shells for stability. Its mode of feeding is uncertain.

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. 1989. Burgess Shale faunas and the Cambrian explosion. Science, 246: 339-346.

CONWAY MORRIS, S. 1993. Ediacaran-like fossils in Cambrian Burgess Shale-type faunas of North America. Palaeontology, 36(3): 593-635.

WALCOTT, C. D. 1911. Middle Cambrian holothurians and medusae. Cambrian Geology and Paleontology II. Smithsonian Miscellaneous Collections, 57(3): 41-68.

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