The Burgess Shale

Acanthotretella spinosa

Reconstruction of Acanthotretella spinosa.

© MARIANNE COLLINS

Taxonomy:

Class: Lingulata (Order: Siphonotretida, stem group brachiopods)
Remarks:

Acanthotretella spinosa is probably related to a primitive group of brachiopods of the Order Siphonotretida (Holmer and Caron, 2006).

Described by: Holmer and Caron
Description date: 2006
Etymology:

Acanthotretella – from the Greek akantha, “thorn,” and tretos, “perforated,” and the Latin diminutive ella, describing the small, perforated, spiny shell.

spinosa – from the Latin spinosus, referring to the exterior spines.

Type Specimens: Holotype –ROM57749 in the Royal Ontario Museum, Toronto, Canada.
Other species:

Burgess Shale and vicinity: none.

Other deposits: Acanthotretella decaius from the early Cambrian Guanshan fauna, China.

Age & Localities:

Species name: Acanthotretella spinosa
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:

Specimens were first illustrated as Lingulella sp. by Jin, et al. (1993), and formally described as Acanthotretella spinosa by Holmer and Caron (2006). New characters preserved in a related species from China (Acanthotretella decaius, Zhifei et al., 2010) reinforce the probable position of this genus within the Order Siphonotretida.

Description:

Morphology:

The shell of Acanthotretella is mainly organic in composition with probably only minor organo-phosphatic mineralization, and is ventri-biconvex. Both valves are covered in long, slender spines that penetrate the shell and are posteriorly inclined, angled obliquely away from the anterior margin. A long, flexible pedicle emerges from an external tube that extends from the pedicle foramen along the ventral valve. The pedicle is at least three to four times the length of the valves. The visceral area of both valves is short and triangular, and does not extend to mid-valve. Other interior features are poorly known.

Abundance:

Most specimens come from the Walcott Quarry and represent one of the rarest brachiopods with less than 0.05% of the entire fauna (Caron and Jackson, 2008).

Maximum Size:
8 mm

Ecology:

Ecological Interpretations:

The long, thin pedicle and overall shell shape probably preclude an infaunal habit. Pedicles of several specimens were found attached at the terminal bulb to organic structures, suggesting that Acanthotretella spinosa was epibenthic. The pedicle was likely able to maintain the shell in an upright position well above the sediment-water interface. Extraction of food particles from the water would have been possible thanks to a 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.

HU, S. X., Z. F. ZHANG, L. E. HOLMER AND C. B. SKOVSTED. 2010. Soft-part preservation in a linguliform brachiopod from the lower Cambrian Wulongqing Formation (Guanshan Fauna) of Yunnan, South China. Acta Palaeontologica Polonica, 55: 495-505.

HOLMER, L. E. AND J.-B. CARON. 2006. A spinose stem-group brachiopod with pedicle from the Middle Cambrian Burgess Shale. Acta Zoologica (Stockholm), 87: 273-290.

JIN, Y. G, X. G. HOU. AND H. Y. WANG. 1993. Lower Cambrian pediculate lingulids from Yunnan, China. Journal of Paleontology, 67: 788-798.

Other Links:

http://onlinelibrary.wiley.com/doi/10.1111/j.1463-6395.2006.00241.x/abstract

Wapkia grandis

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

ANIMATION BY PHLESCH BUBBLE © ROYAL ONTARIO MUSEUM

Taxonomy:

Class: Demospongia (Order: Monaxonida)
Remarks:

Wapkia 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: Wapkia grandis
Described by: Walcott
Description date: 1920
Etymology:

Wapkia – origin of name is unknown

grandis – from the Latin grandis, “large.” This name refers to the large size and complex skeleton of this sponge.

Type Specimens: Lectotype –USNM66458 (W. grandis), in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA. Holotype –ROM53544 (W. elongata), in the Royal Ontario Museum, Toronto, Canada.
Other species:

Burgess Shale and vicinity: W. elongata Rigby and Collins, 2004 from the Tulip Beds (S7) on Mount Stephen.

Other deposits: none.

Age & Localities:

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

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

History of Research:

Brief history of research:

Wapkia was described by Walcott in his initial description of the Burgess Shale sponges in 1920. The genus was re-examined by Rigby in 1986. Rigby and Collins (2004) also redescribed the genus and proposed a new species, W. elongata.

Description:

Morphology:

Wapkia is a large elongate or oval sponge with bundles of coarse and fine spicules aligned in long vertical columns and distinct horizontal bundles. The surface of the sponge is smooth and lacks any vertical or horizontal ridges. Spicules are straight and pointed at both ends (oxeas). The exact position of the various bundles of spicules in the skeleton is still uncertain, but it seems that the inner part of the skeleton is reticulate with horizontal wrinkles that are typical of the species and produced by horizontal bundles of spicules. The dermal layer is formed by bundles of oxeas up to 60 mm long which give a characteristic plumose aspect to this sponge. W. elongata is distinguished from W. grandis based on the overall shape of the sponge and different skeletal structures (varying distance between the horizontal spicule bundles).

Abundance:

Wapkia is rare and represents only 0.06% of the Walcott Quarry community (Caron and Jackson, 2008).

Maximum Size:
170 mm

Ecology:

Ecological Interpretations:

Wapkia 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. 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.

Other Links:

None

Walcottidiscus typicalis

Walcottidiscus typicalis (GSC 45368). Complete but poorly preserved specimen. Specimen diameter = 18 mm. Specimen dry – direct light. Walcott Quarry.

© GEOLOGICAL SURVEY OF CANADA. PHOTO: JEAN-BERNARD CARON

Taxonomy:

Class: Edrioasteroidea (Order: Edrioasteroida, stem group echinoderms)
Remarks:

Walcottidiscus is a poorly known edrioasteroid, an extinct group of echinoderms (Smith, 1985).

Species name: Walcottidiscus typicalis
Described by: Bassler
Description date: 1935
Etymology:

Walcottidiscus – from Charles Walcott, the discoverer of the Burgess Shale, and the Greek diskos, “disc.” The name refers to the flattened appearance of the fossils.

typicalis – from the Greek typikos, “type,” perhaps in reference to the single specimen originally described.

Type Specimens: Holotype –USNM90754 (W. typicalis),USNM90755 (W. magister) in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.
Other species:

Burgess Shale and vicinity: W. magister Bassler, 1935 from the Walcott Quarry on Fossil Ridge (but see below paragraph brief history of research).

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 species known from a single specimen each were originally described by Bassler in 1935 and 1936: a small form W. typicalis, and a larger form W. magister respectively. However, W. magister is now thought to belong to W. typicalis (Smith, 1985) but additional fossil material would be required to confirm this hypothesis. Walcottidiscus resembles Kailidiscus chinensis, a chinese form from the Middle Cambrian Kaili deposit, but remains too poorly known to draw more detailed comparisons between the two genera (Zhao et al., 2010).

Description:

Morphology:

The body (theca) is ovoid in outline and has a relatively small dorsal surface compared to the ventral one. The upper central part of the theca is not calcified, but the outer zone is composed of small calcified plates. A five star-shaped food groove lined with small plates (the ambulacra) is present on the upper surface. The five arms of the ambulacra are arranged in a 2:1:2 fashion around the mouth, and they are at first straight and then turn to the left near the edge of the theca. Differences between the two species are the size and degree of ambulacral curvature, but those differences could simply be a factor of growth.

Abundance:

Walcottidiscus is very rare only two specimens were originally described. A few additional specimens are known in the Burgess Shale collections of the Geological Survey of Canada and the Royal Ontario Museum.

Maximum Size:
64 mm

Ecology:

Ecological Interpretations:

Walcottidiscus was most likely resting on the seafloor. Food particles were transported by food grooves (ambulacrum) into a central mouth at the top of the theca.

References:

BASSLER, R. S. 1935. The classification of the Edrioasteroidea. Smithsonian Miscellaneous Collections, 93: 1-11.

BASSLER, R. S. 1936. New species of American Edrioasteroidea. Smithsonian Miscellaneous Collections, 95: 1-33.

SMITH, A. B. 1985. Cambrian eleutherozoan echinoderms and the early diversification of edrioasteroids. Palaeontology, 28: 715-756.

ZHAO, Y., C. D. SUMRALL, R. L. PARSLEY AND J. I. N. PENG. 2010. Kailidiscus, a new plesiomorphic edrioasteroid from the basal Middle Cambrian Kaili biota of Guizhou province, China. Journal of Paleontology, 84: 668-680.

Other Links:

None

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.

Other Links:

None

Ulospongiella ancyla

Ulospongiella ancyla (ROM 43830) – Holotype. Nearly complete individual. Specimen height = 19 mm. Specimen dry – direct light (left), wet – polarized light (right). Trilobite Beds on Mount Stephen.

© ROYAL ONTARIO MUSEUM. PHOTOS: JEAN-BERNARD CARON

Taxonomy:

Class: Demospongea (Order: Monaxonida)
Remarks:

Ulospongiella 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: Ulospongiella ancyla
Described by: Rigby and Collins
Description date: 2004
Etymology:

Ulospongiella – from the Greek oulus, “wooly or curly,” and spongia, “sponge.” The name refers to the curled or curved spicules forming the skeleton.

ancyla – from the Greek anklyos, “bent or hooked.” The name makes reference to the curved spicules.

Type Specimens: Holotype –ROM43830 (wrongly referred asROM48830 in Rigby and Collins 2004) 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 Trilobite Beds on Mount Stephen.

History of Research:

Brief history of research:

Ulospongiella was described by Rigby and Collins in 2004 based on collections made by the Royal Ontario Museum.

Description:

Morphology:

Ulospongiella is a small sponge less than 2 cm in height. Its shape is subcyclindrical with a rounded base. Most spicules forming the skeleton are pointed at both ends (oxeas). These oxeas are strongly curved or hooked shape and form a relatively dense mesh. A few coarser and longer spicules with a round base extend upward from the wall. There is no clear indication of canals within the sponge there is no evidence of a central cavity (spongocoel).

Abundance:

Only three specimens are known, all from the Trilobite Beds.

Maximum Size:
19 mm

Ecology:

Ecological Interpretations:

Ulospongiella 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:

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.

Other Links:

None

Tubulella flagellum

Tubulella flagellum (ROM 59942) – Proposed Lectotype. Figures 1a of Matthew (1899) and photograph of original specimen (right). Approximate specimen length = 80 mm. Specimen dry – direct light. Trilobite Beds on Mount Stephen.

© ROYAL ONTARIO MUSEUM. PHOTOS: JEAN-BERNARD CARON

Taxonomy:

Class: Unranked clade (stem group cnidarians)
Remarks:

This fossil was originally thought to represent the tube of some sedentary polychaete worms (Matthew, 1899; Howell, 1949), but has more recently been compared to the sessile polyp stage of a scyphozoan jellyfish that builds tapered, chitinous tubes fixed to the substrate by an attachment disc (Van Iten et al., 2002).

Species name: Tubulella flagellum
Described by: Matthew
Description date: 1899
Etymology:

Tubulella – from the latin tubulus, “tube, or tubule,” and the suffix –ella, denoting “little.”

flagellum – the Latin for “whip,” in allusion to the long, tapering form of the tubular theca.

Type Specimens: Syntype–ROM59942 in the Royal Ontario Museum, Toronto, ON, Canada.
Other species:

Burgess Shale and vicinity: Many shared similarities suggest that other thecate Burgess Shale fossils such as Byronia annulataSphenothallus sp., Cambrorhytium major, and Cfragilis may be related to Tubulella.

Other deposits: Other species occur worldwide in rocks from the Cambrian period.

Age & Localities:

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

The Trilobite Beds, Tulip Beds (S7) and additional smaller localities on Mount Stephen. The Walcott and Raymond Quarries on Fossil Ridge, Mount Odaray and Monarch Cirque.

History of Research:

Brief history of research:

In August 1887 the Toronto meeting of the British Association for the Advancement of Science was followed by a special geological rail tour to western Canada organized by Byron Edmund Walker (a prominent Canadian banker). One of the excursion highlights was a visit to the Mount Stephen Trilobite Beds, after which Walker loaned his personal collection of Mount Stephen fossils to Canada’s leading Cambrian palaeontologist, George F. Matthew, of Saint John, New Brunswick. In 1899, Matthew published a series of new descriptions based on this material, including Urotheca flagellum, a rare form he interpreted as whip-shaped worm tube, illustrated in two engravings. Walker donated these fossils to the University of Toronto in 1904, and in 1913 they were transferred to the new Royal Ontario Museum of Palaeontology. In 1949, American palaeontologist B. F. Howell found that Matthew’s genus name Urotheca was already in use for a living reptile, so he substituted it for the new name Tubulella. Subsequently, this and similar fossils were reinterpreted as cnidarian polyp thecae. The single best specimen of Walker’s Urotheca flagellum remained unrecognized until it was “rediscovered” in the ROM collections in 2010.

Description:

Morphology:

The chitinous or chitinophosphatic tube (theca) of Tubulella flagellum is a very long and slender cone, with a maximum diameter of about 4 mm. The thecae may be almost straight, or show varying degrees of curvature. The thecal wall is relatively thick and often appears densely black against the shale matrix. The external surface shows very fine transverse growth lines, but usually no strong annular ridges. Often, two or more lengthwise creases or ridges were formed as the result of the crushing and compaction of the tube’s original circular or oval cross section. Some specimens possess a combination of features seen in Tubulella and Byronia, with very narrow thecae bearing both annulae and longitudinal creases. Small clusters of such Tubulella-like thecae are occasionally found closely associated with Byronia annulata, but it is not known whether these were asexually generated “buds” or discrete organisms growing attached to the larger tubes. No soft tissues of Tubulella flagellum have been described to date.

Abundance:

Uncommon in the Trilobite Beds on Mount Stephen. Relatively common in the Walcott Quarry on Fossil Ridge where it represents about 0.25% of the specimens in the community (Caron and Jackson, 2008).

Maximum Size:
100 mm

Ecology:

Ecological Interpretations:

The theca of Tubulella was likely attached to the substrate using an apical disc which is usually broken off. The absence of soft tissue preservation makes the assignment to a particular feeding strategy tentative. By comparison with forms such as Cambrorhytium, a carnivorous or suspension feeding habit seems possible.

References:

BISCHOFF, C. O. 1989. Byroniida new order from early Palaeozoic strata of eastern Australia (Cnidaria, thecate scyphopolyps). Senkenbergiana Lethaea, 69(5/6): 467-521.

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 R. A. ROBISON. 1988. More soft-bodied animals and algae from the Middle Cambrian of Utah and British Columbia. The University of Kansas Paleontological Contributions, Paper 122: 48 pp.

HOWELL, B. F. 1949. New hydrozoan and brachiopod and new genus of worms from the Ordovician Schenectady Formation of New York. Bulletin of the Wagner Free Institute of Science, 24(1): 8 pp.

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.

RASETTI, F. 1951. Middle Cambrian stratigraphy and faunas of the Canadian Rocky Mountains. Smithsonian Miscellaneous Collections, 116(5): 277 pp.

VAN ITEN, H., M.-Y. MAO-YAN, AND D.COLLINS 2002. First report of Sphenothallus Hall, 1847 in the Middle Cambrian. Journal of Paleontology, 76: 902-905.

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

ZHU, M.-Y., H. VAN ITEN, R. S. COX, Y.-L. ZHAO AND B.-D. ERDTMANN. 2000. Occurrence of Byronia Matthew and Sphenothallus Hall in the Lower Cambrian of China. Paläontologische Zeitschrift, 74: 227-238.

Other Links:

None

Thaumaptilon walcotti

Thaumaptilon walcotti (USNM 468028) – Holotype, part and counterpart. Complete specimen. Specimen height = 212 mm. Specimen dry – direct light (far left and far right), wet – polarized light (middle images). Walcott Quarry.

© SMITHSONIAN INSTITUTION – NATIONAL MUSEUM OF NATURAL HISTORY. PHOTOS: JEAN-BERNARD CARON

Taxonomy:

Class: Unranked clade (stem group cnidarians)
Remarks:

Thaumaptilon was first interpreted as a Cambrian member of the frondose Ediacaran Biota, related to cnidarians and particularly to a group of modern anthozoans called pennatulaceans or sea pens (Conway Morris, 1993). This connection is no longer widely accepted (Antcliffe and Brasier, 2008); Thaumaptilon has also been proposed as a critical link between Ediacaran fronds and ctenophores (Dzik, 2002). A position in the cnidarian stem group (i.e. more primitive than the anthozoans) has been supported by the discovery of similar fossils in the Chengjiang Biota (Shu et al., 2006).

Species name: Thaumaptilon walcotti
Described by: Conway Morris
Description date: 1993
Etymology:

Thaumaptilon – from the Greek thauma, “wonderful,” and ptilon, “soft feather,” after its feather-like appearance.

walcotti – after Charles Walcott, discoverer of the Burgess Shale.

Type Specimens: Holotype –USNM468028 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.

History of Research:

Brief history of research:

Walcott had studied and photographed Thaumaptilon, but never published his work. The fossil specimens were formally described in 1993 by Conway Morris, who had also alluded to them in previous work (1979; 1989; 1990).

Description:

Morphology:

Thaumaptilon is an oblong frond that somewhat resembles a feather; it is bilaterally symmetrical, with a central axis supporting a number of lateral branches. The branches appear to be connected to one another by narrow canals. A blunt holdfast attached the animal to the sea floor. Of the three known specimens, the largest is 21 cm tall and reaches 5 cm across; the smaller specimens – presumed to be juveniles – are only a few centimetres long. The frond is flattened, and tapers slightly towards its tip. It consists of about three dozen branches angled at 45º to the central axis, and primarily grew by inflation – perhaps with some addition of branches by apical budding. Unlike modern sea pens, Thaumaptilon’s branches attach to a common base. Lines of pustules on one side of the frond have been interpreted as retracted zooids (individual members of a colonial organism), which are arranged very haphazardly in comparison to the neat combs seen in modern sea pens.

Abundance:

Only three specimens are known.

Maximum Size:
210 mm

Ecology:

Ecological Interpretations:

The holdfast would have anchored the organism to the soft sediment of the sea floor, and could perhaps contract to adjust the height and angle of the frond. Based on the interpretation of the pustules as zooids, a colonial, suspension-feeding lifestyle has been proposed. It has been suggested that Thaumaptilon could retract into its stem when threatened, for protection (Conway Morris, 1998).

References:

ANTCLIFFE, J. B. AND M. D. BRASIER. 2008. Charnia at 50: Developmental models for Ediacaran fronds. Palaeontology, 51(1): 11-26.

CONWAY MORRIS, S. 1979. The Burgess Shale (Middle Cambrian) fauna. Annual Review of Ecology and Systematics, 10(1): 327-349.

CONWAY MORRIS, S. 1989. Burgess Shale faunas and the Cambrian explosion. Science, 246(4928): 339.

CONWAY MORRIS, S. 1990. Late Precambrian and Cambrian soft-bodied faunas. Annual Review of Earth and Planetary Sciences, 18(1): 101-122.

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

CONWAY MORRIS, S. 1998. The Crucible of Creation, the Burgess Shale and the Rise of Animals. Oxford University Press, 242 p.

SHU, D. G., S. CONWAY MORRIS, J. HAN, Y. LI, X. L. ZHANG, H. HUA, Z. F. ZHANG, J. N. LIU, J. F. GUO, Y. YAO AND K. YASUI. 2006. Lower Cambrian vendobionts from China and early diploblast evolution. Science, 312(5774): 731-734.

Other Links:

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

Stephenospongia magnipora

Stephenospongia magnipora (ROM 43127) – Holotype. Fragment of the only known specimen of the species showing large holes in the wall of this sponge. Specimen height = 44 mm. Specimen dry – polarized light. Trilobite Beds on Mount Stephen.

© ROYAL ONTARIO MUSEUM. PHOTO: JEAN-BERNARD CARON

Taxonomy:

Class: Hexactinellida (Order: Reticulosa)
Remarks:

Stephenospongia is placed in the Family Hintzespongiidae (primitive hexactinellids). Hexactinellid sponges (glass sponges) have a skeleton composed of four to six-pointed spicules. They are considered to be an early branch within the Porifera phylum due to their distinctive composition.

Species name: Stephenospongia magnipora
Described by: Rigby
Description date: 1986
Etymology:

Stephenospongia – from Mount Stephen (3,199 m), a mountain peak in Yoho National Park, named after George Stephen (1829 – 1921), first president of the Canadian Pacific Railway and the Latin spongia, meaning “sponge.”

magnipora – from the Latin magnus, “great,” and porus, “pore.” The name makes reference to the large pores present in the skeleton of this sponge.

Type Specimens: Holotype –ROM43127 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 Trilobite Beds on Mount Stephen.

History of Research:

Brief history of research:

Stephenospongia was described by Rigby (1986) (see also Rigby and Collins 2004) based on a single specimen discovered by the Royal Ontario Museum in 1982.

Description:

Morphology:

Stephenospongia has a conical and almost cylindrical shape. The skeleton is composed of six rayed spicules (called hexactines) typical of the hexactinellid sponges. The spicules mesh together to form a single layer and are arranged in an irregular fashion especially around holes in the sponge wall. Prominent holes organized in vertical and horizontal rows are separated by tracts of spicules with ray lengths reaching more than one centimetre. The basal and top parts are not preserved.

Abundance:

Only a single specimen is known and comes from the Trilobite Beds.

Maximum Size:
44 mm

Ecology:

Ecological Interpretations:

Stephenospongia 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:

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.

Other Links:

None

Sphenothallus sp.

Taxonomy:

Sphenothallus sp. (GSC 134789). Fragment of a large specimen showing longitudinal thickenings clearly differentiated near the aperture area (to the right). A Micromitra (Dictyonina) brachiopod is attached to the lower part of the tube. Approximate specimen length = 50 mm. Specimen dry – direct light. Trilobite Beds on Mount Stephen.

© GEOLOGICAL SURVEY OF CANADA. PHOTO: JEAN-BERNARD CARON

Class: Unranked clade (stem group cnidarians)
Remarks:

Sphenothallus has been compared to some form of tubiculous annelid worm or the sessile polyp stage of a scyphozoan jellyfish that builds tapered, chitinous tubes fixed to the substrate by an attachment disc (Van Iten et al., 2002).

Species name: Sphenothallus sp.
Described by: Van Iten et al.
Description date: 2002
Etymology:

Sphenothallus – from the Greek sphen, “wedge”, and thallos, “branch.”

Species name not determined.

Type Specimens: Not applicable
Other species:

Burgess Shale and vicinity: Many shared similarities suggest that other thecate Burgess Shale fossils such as Byronia annulataCambrorhytium majorCfragilis and Tubulella flagellum, may be related to Sphenothallus sp.

Other deposits: Other species occur worldwide in rocks from the Cambrian to the Silurian periods. Sphenothallus is also known in the Kaili Formation (Zhu et al., 2000).

Age & Localities:

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

The Trilobite Beds on Mount Stephen.

History of Research:

Brief history of research:

Two specimens from the Trilobite Beds were illustrated in 2002 (Van Iten et al.). A third previously unrecognized specimen was identified in the Geological Survey of Canada collections in Ottawa (Billings collection) in the Spring of 2010. Owing to the relatively low degree of morphological variations among all known species, it is not currently possible to assign the Burgess Shale form to any particular species without better preserved specimens.

Description:

Morphology:

The chitinophosphatic tube (theca) of Sphenothallus consists of longitudinal thickenings which are particularly obvious towards the aperture area. The tube is gently curved and does not seem to branch. The maximum diameter of the largest specimen is about 4 mm for a length of about 75 mm. A thin wall is present between the longitudinal thickenings and terminates in a smooth margin near the aperture, a couple of millimeters beyond the longitudinal thickenings. The tube is roughly circular in the apical region and is very slender, with the two longitudinal thickenings less differentiated in this area. The surface of the entire tube including thickenings is smooth with no evidence of ridges or annulations. All three specimens lack the apical ends, so it is not evident that this species had a holdfast and there is no evidence of soft-tissue preservation.

Abundance:

Only three specimens known from the Trilobite Beds on Mount Stephen.

Maximum Size:
75 mm

Ecology:

Ecological Interpretations:

The theca of Sphenothallus was likely attached to the substrate via an apical disc as can be seen in other better known species. The absence of soft tissue preservation makes the assignment to a particular feeding strategy tentative. By comparison with possible related forms such as Cambrorhytium, a carnivorous or suspension feeding habit seems possible.

References:

VAN ITEN, H., M.-Y. ZHU AND D. COLLINS. 2002. First report of Sphenothallus Hall, 1847 in the Middle Cambrian. Journal of Paleontology, 76: 902-905.

ZHU, M.-Y., H. VAN ITEN, R. S. COX, Y.-L. ZHAO AND B.-D. ERDTMANN. 2000. Occurrence of Byronia Matthew and Sphenothallus Hall in the Lower Cambrian of China. Paläontologische Zeitschrift, 74: 227-238.

Other Links:

None

Sarotrocercus oblita

Reconstruction of Sarotrocercus oblita.

© MARIANNE COLLINS

Taxonomy:

Class: Unranked clade (stem group arthropods)
Remarks:

The phylogenetic affinity of Sarotrocercus is uncertain because its morphology is too poorly known to make a definitive designation. Fryer (1998) suggested it was the most primitive of all arthropods, and it was placed within the Arachnomorpha by Cotton and Braddy (2004). Sarotrocercus has also been aligned with Megacheiran taxa such as Yohoia (e.g. Briggs and Fortey, 1989) and Leanchoilia (e.g., Wills et al. 1995; 1998).

Species name: Sarotrocercus oblita
Described by: Whittington
Description date: 1981
Etymology:

Sarotrocercus – from the Greek sarotes, “sweeper”, and kerkops, “a long tailed-monkey”, in reference to the feathery aspect of the tail.

oblita – from the Latin oblitus, “forgotten”, perhaps in reference to the fact that the few specimens of this species were described as part of another species.

Type Specimens: Holotype –USNM144890 (part) and UNSM 272171 (counterpart) 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.

History of Research:

Brief history of research:

The genus Sarotrocercus was erected by Harry Whittington in 1981 based on seven specimens originally included within Molaria spinifera (Simonetta and Delle Cave, 1975). No further research has been performed on the fossil material since then, although Sarotrocercus has been included in many studies of arthropod relationships (e.g. Briggs and Fortey, 1989; Wills et al., 1995; Fryer, 1998).

Description:

Morphology:

Sarotrocercus has an oval body consisting of a head shield and nine overlapping trunk segments; a cylindrical posterior segment carries a relatively short, narrow spine ending in a fan-shape cluster of small spikes. The whole animal was about 1.5 cm long. Although the head shield was not very strongly developed, it did bear a pair of large, stalked eyes that poked out from beneath the margin, and a pair of jointed appendages. Each of the nine body segments bore a pair of lobate appendages, with comb-like fringes which might have functioned as gills.

Abundance:

S. oblita is rare in the Burgess Shale. It was originally described on the basis of 7 specimens (Whittington, 1981), and 28 further specimens have been recovered from the Walcott Quarry representing less than 0.1% of the community (Caron and Jackson, 2008).

Maximum Size:
16 mm

Ecology:

Ecological Interpretations:

The absence of walking limbs combined with an inferred flexibility of the body imply that the organism swam, probably in an inverted position, using its paddle-like appendages and long tail. Its rarity in the Burgess Shale suggests that it may have spent much time in the water column, thus avoiding submarine landslides that trapped animals living on the sea floor. The absence of sediment in its gut suggest that Sarotrocercus was a filter feeder (Briggs and Whittington, 1985; Whittington, 1981).

References:

BRIGGS, D. E. G. AND R. A. FORTEY, 1989. The Early radiation and relationships of the major arthropod groups. Science, 246: 241-243.

BRIGGS, D. E. G. AND H. B. WHITTINGTON, 1985. Modes of life of arthropods from the Burgess Shale, British Columbia. Transactions of the Royal Society of Edinburgh. Earth Sciences, 76(2-3): 149-160.

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

COTTON, T. J. AND S. J. BRADDY, 2004. The phylogeny of arachnomorph arthropods and the origin of the Chelicerata. Transactions of the Royal Society of Edinburgh, 94(03): 169-193.

FRYER, G. 1998. A defence of arthropod polyphyly, p. 23. In R. A. Fortey and R. H. Thomas (eds.), Arthropod relationships. Springer, London.

SIMONETTA, A. M. AND L. DELLE CAVE, 1975. The Cambrian non-trilobite arthropods from the Burgess shale of British Columbia: A study of their comparative morphology, taxonomy and evolutionary significance. Palaeontographia Italica, 69: 1-37.

WHITTINGTON, H. B. 1981. Rare arthropods from the Burgess Shale, Middle Cambrian, British Columbia. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences, 292(1060): 329-357.

WILLS, M. A., D. E. G. BRIGGS, R. A. FORTEY AND M. WILKINSON, 1995. The significance of fossils in understanding arthropod evolution. Verhandlungen den deutschen zoologischen Gesellschaft, 88: 203-216.

WILLS, M. A., D. E. G. BRIGGS, R. A. FORTEY, M. WILKINSON AND P. H. A. SNEATH, 1998. An arthropod phylogeny based on fossil and recent taxa, p. 33-105. In G. D. Edgecombe (ed.), Arthropod fossils and phylogeny. Columbia University Press, New York.

Other Links:

None