The Burgess Shale

Life Habits: Epibenthic

Protoprisma annulata

Protoprisma annulata (ROM 53557) – Holotype. Nearly complete specimen showing a clump of branches attached to a basal part (coated with ammonium chloride sublimate to show details). Specimen height = 150 mm. Specimen dry – direct light. Tulip Beds (S7) on Mount Stephen.

© Royal Ontario Museum. Photo: Jean-Bernard Caron

Taxonomy:

Kingdom: Epibenthic
Phylum: Epibenthic
Higher Taxonomic assignment: Hexactinellida (Order: Reticulosa)
Species name: Protoprisma annulata
Remarks:

Hexactinellid sponges (glass sponges) have a skeleton composed of four to six-pointed siliceous spicules. They are considered to be an early branch within the Porifera phylum due to their distinctive composition.

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

Protoprisma – from the Greek protos, “first,” and prisma, “prism.” This name refers to the early occurrence of this prismatic sponge.

annulata – from the Latin annulatus, meaning “ringed, or circular.” The name makes reference to the annulated growth form of this species.

Type Specimens: Holotype –ROM53557, in the Royal Ontario Museum, Toronto, Canada.
Other species:

Burgess Shale and vicinity: none.

Other deposits: none.

Age & Localities:

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

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

History of Research:

Brief history of research:

Ribgy and Collins described this genus in 2004 based on material collected by the Royal Ontario Museum.

Description:

Morphology:

This sponge has an elongate annulated shape with several branches, which give it a hand-like appearance. Each branch has vertical angular ridges which results in a prismatic cross section. The ridges and the troughs between them are composed of fine hexactine spicules, cross-connected by horizontal strands that thatch the skeleton together. The type specimen is almost complete at 15 cm tall and shows that all of the branches originate from a central point at the base. The base of the sponge would have had an attachment structure to keep the sponge anchored in the sediment surface. As neither of the two specimens recovered are complete, it is not known what the top of this sponge would have looked like.

Abundance:

Protoprisma is known only from two specimens, one collected from the Tulip Bed (S7) locality on Mount Stephen and one from the Raymond Quarry on Fossil Ridge.

Maximum Size:
150 mm

Ecology:

Life habits: Epibenthic
Feeding strategies: Epibenthic
Ecological Interpretations:

Protoprisma 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

Priscansermarinus barnetti

Priscansermarinus barnetti (ROM 36064) – Holotype (specimen A). Slab with 62 individuals including the holotype (white frame to the left and close up to the right). Specimen length (holotype) = 32 mm. Specimen dry – polarized light (both images). Walcott Quarry talus.

© Royal Ontario Museum. Photos: Jean-Bernard Caron

Taxonomy:

Kingdom: Epibenthic
Phylum: Epibenthic
Higher Taxonomic assignment: Non applicable
Species name: Priscansermarinus barnetti
Remarks:

Originally proposed as a crustacean arthropod and a possible member of the pedunculate lepadomorph barnacles within Maxillopoda (Subclass Thecostraca) (Collins and Rudkin, 1981), its affinities have since been questioned and remain equivocal.

Described by: Collins and Rudkin
Description date: 1981
Etymology:

Priscansermarinus – from the Latin priscus “of ancient times;” anser, “goose” and marinus, “sea,” (together forming “sea goose”) in reference to the modern goose barnacles.

barnetti – after Robert Barnett, member of the 1975 ROMexpedition, who found the first specimens.

Type Specimens: Holotype –ROM36064a in the Royal Ontario Museum, Toronto, Ontario, Canada
Other species:

Burgess Shale and vicinity: none.

Other deposits: none.

Age & Localities:

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

The Walcott and Raymond Quarries on Fossil Ridge, Mount Field.

History of Research:

Brief history of research:

The “discovery” slab bearing 62 individuals of a previously unknown organism was found by Robert Barnett in talus (scree) beneath the Walcott Quarry level during the ROM’s inaugural Burgess Shale expedition in 1975. Priscansermarinus barnetti was described and named in 1981, and interpreted as a probable stalked (pedunculate) lepadomorph (goose) barnacle, pending recovery of additional specimens preserving definitive characters. The barnacle, and even the arthropod, affinities of Priscansermarinus have since been questioned (Briggs, 1983; Briggs et al., 2005).

Description:

Morphology:

Priscansermarinus consists of two primary components – an ovate triangular shaped, laterally compressed “body,” and a short, thick “stalk.” The body region shows a highly reflective centralized subtriangular region that was originally interpreted as evidence for a thin non-biomineralized external “plate” on either side of the body. This is now recognized to be an internal structure of greater anatomical complexity. The stalk, or stolon, which appears to emerge from the body rather than blend into it, is cylindrical in shape and at least moderately flexible; the distal end bears a terminal disc exhibiting a radiating pattern. In most known specimens, the stalk comprises slightly more than half of the total length of the animal.

Abundance:

Moderately common at some Raymond Quarry levels; uncommon elsewhere.

Maximum Size:
50 mm

Ecology:

Life habits: Epibenthic
Feeding strategies: Epibenthic
Ecological Interpretations:

This species is too poorly known to describe its ecology with great certainty. The terminal disk at the base of its stolon was probably used for anchoring the animal in or on the mud. Without any apparent tentacles and obvious feeding structures, a suspension feeding mode of life is a strong possibility.

References:

BRIGGS, D. E. G. 1983. Affinities and early evolution of the Crustacea: the evidence of the Cambrian fossils, p. 1-22. In F. R. Schram (ed.), Crustacean Phylogeny. Balkema, Rotterdam.

BRIGGS, D. E. G., M. D. SUTTON AND D. J. SIVETER. 2005. Metamorphosis in a Silurian barnacle. Proceedings of the Royal Society, B, 272: 2365-2369.

COLLINS, D. AND RUDKIN, D. M. 1981. Priscansermarinus barnetti, a probable lepadomorph barnacle from the Middle Cambrian Burgess Shale of British Columbia. Journal of Paleontology, 55: 1006-1015.

Other Links:

None

Leptomitus lineatus

Leptomitus undulatus (ROM 53571) – Holotype (part and counterpart). Only known specimen of this species showing partial base, prominent ridges and top part (osculum). Specimen height = 78 mm. Specimen wet – direct light. Walcott Quarry.

© Royal Ontario Museum. Photos: Jean-Bernard Caron

Taxonomy:

Kingdom: Epibenthic
Phylum: Epibenthic
Higher Taxonomic assignment: Demospongea (Order: Monaxonida)
Species name: Leptomitus lineatus
Remarks:

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

Described by: Walcott
Description date: 1920
Etymology:

Leptomitus – from the Greek lept, “slender,” and mitos, “thread.” This name refers to the overall shape of the sponge.

lineatus – from the Latin lineatus, “streaked.” This refers to the wrinkle appearance of this sponge.

Type Specimens: Lectotype –USNM66448 (L. lineatus) in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA. Holotype –ROM53558 (L. undulatus) in the Royal Ontario Museum, Toronto, Canada.
Other species:

Burgess Shale and vicinity: L. undulatus Rigby and Collins 2004 from the Walcott Quarry.

Other deposits: L. zitteli Walcott, 1886 from the Middle Cambrian Parker Slate in Vermont; L. metta Rigby, 1983 from the Middle Cambrian Marjum Formation of Utah; L. conicus García-Bellido et al., 2007 from the Middle Cambrian Murero Formation of Spain; L. teretiusculus Chen, Hou and Lu, 1989 from the Lower Cambrian Chengjiang biota in China (see Rigby and Hou, 1995); unidentified species from the Lower Cambrian Niutitang Formation in China (Yang et al., 2003).

Age & Localities:

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

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

History of Research:

Brief history of research:

Leptomitus was originally described by Charles Walcott (1920) as a new genus “Tuponia” along with several species (T. lineatea, T. flexilis, T. flexilis var. intermedia). This genus was later synonymized by Resser and Howell (1938) with Leptomitus, a genus named by Walcott in 1886. Ribgy (1986) redescribed the Burgess Shale sponges including Leptomitus and considered L. flexilis to be a junior synonym of L. lineatus. Rigby and Collins (2004) added a second species L. undulatus based on new material collected by the Royal Ontario Museum.

Description:

Morphology:

L. lineatus is an elongate tubular sponge with a double-layered skeleton. The outer layer is composed of long monoaxial spicules (simple spicules with pointed ends) arranged vertically along the length of the sponge. The varying thicknesses of these elongate spicules give the sponge a distinctive wrinkly appearance in the fossils. The inner layer is composed of tiny horizontal spicules that form an unclumped thatch; these tufts can be seen at the oscular margin (opening at the top of the sponge). The base of the sponge is rounded in shape and would have had a small holdfast structure. L. undulatus has the same wall structure as L. lineatus but has a rounder goblet shaped skeleton.

Abundance:

L. lineatus is relatively common in the Walcott Quarry and represents 0.26% of the community (Caron and Jackson, 2008). L. undulatus is known from a single specimen.

Maximum Size:
360 mm

Ecology:

Life habits: Epibenthic
Feeding strategies: Epibenthic
Ecological Interpretations:

Leptomitus 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.

CHEN, J. Y., X. G. HOU AND H. Z. LU. 1989. Lower Cambrian leptomitids (Demospongea), Chengjiang, Yunnan. Acta Palaeontologica Sinica, 28: 17-31.

GARCÍA-BELLIDO, D. C., R. GOZALO, J. B. CHIRIVELLA MARTORELL AND E. LIÑÁN. 2007. The demosponge genus Leptomitus and a new species from the Middle Cambrian of Spain. . Palaeontology, 50: 467-478.

RESSER, C. F. AND B. F. HOWELL. 1938. Lower Cambrian Olenellus Zone of the Appalachians. Geological Society of American Bulletin, 49: 195-248.

RIGBY, J. K. 1983. Sponges of the Middle Cambrian Marjum Limestone from the House Range and Drum Mountains of Western Millard County, Utah. Journal of Paleontology, 57: 240-270.

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.

YANG, R., W. ZHANG, L. JIANG AND H. GAO. 2003. Chengjiang biota from the Lower Cambrian Niutitang Formation, Zunyi County, Guizhou Province, China. Acta Palaeontologica Sinica, 77: 145-150.

WALCOTT, C. 1886. Second contribution to the studies on the Cambrian faunas of North America. U.S. Geological Survey Bulletin, 30: 1-369.

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

Other Links:

None

Hazelia palmata

3D animation of Hazelia conferta and other sponges (Choia ridleyi, Diagoniella cyathiformis, Eiffelia globosa, Pirania muricata, 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:

Kingdom: Epibenthic
Phylum: Epibenthic
Higher Taxonomic assignment: Demospongea (Order: Monaxonida)
Species name: Hazelia palmata
Remarks:

Hazelia is considered a primitive demosponge, close to Falospongia and Crumillospongia (Rigby, 1986). Demosponges, the same group that are harvested as bath sponges, represent the largest class of sponges today.

Described by: Walcott
Description date: 1920
Etymology:

Hazelia – from Hazel Peak (3,151 m), the older name for Mount Aberdeen, located 4 km SSW of Lake Louise in Banff National Park, Alberta. Mount Aberdeen was named in honor of Lord Gordon in 1897, the Marquis of Aberdeen and the Governor General of Canada from 1893 to 1898.

palmata – from the Latin palm, “palm of the hand,” referring to the broad cup-shape of this sponge and its resemblance to a cupped hand.

Type Specimens: Lectotypes – USNM 66463 (H. palmata – type species), 66465 (H. delicatula), USNM 66505 (H. dignata), USNM 66473 (H. grandis), USNM 66474 (H. nodulifera), USNM 66472 (H. obscura); Holotypes – USNM 66476 (H. conferta), USNM 66779 (H. crateria), USNM 66475 (H. luteria) in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA. Holotype –ROM53573 (H. lobata) in the Royal Ontario Museum, Toronto, Canada.
Other species:

Burgess Shale and vicinity: H. conferta Walcott, 1920, H. crateria Rigby, 1986, H. delicatula Walcott, 1920, H. dignata Walcott, 1920, H. grandis Walcott, 1920, H. lobata Rigby and Collins, 2004, H. luteria Rigby, 1986, H. nodulifera Walcott, 1920, H. obscura Walcott, 1920. Most species known from the Walcott Quarry (See Rigby, 1986 and Rigby and Collins, 2004).

Other deposits: H. walcotti (Resser and Howell, 1938) from the Early Cambrian Kinzers Formation of Pennsylvania (See Rigby, 1987).

Age & Localities:

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

Burgess Shale and vicinity: Hazelia is particularly common in the Walcott Quarry and is less common in the Raymond and Collins Quarries on Fossil Ridge. Many species also occur on Mount Stephen at the Trilobite Beds, Tulip Beds (S7), and other smaller localities.

Other deposits: H. palmata Walcott, 1920 from the Middle Cambrian Marjum Formation (Rigby et al., 1997).

History of Research:

Brief history of research:

Walcott described seven species of Hazelia in his 1920 paper on the Burgess Shale sponges. The genus was redescribed by Rigby in 1986 when two new species were added and one excluded from the genus (H. mammillata now referred to Moleculospina mammillata). Rigby and Collins (2004) added another species based on new material collected by the Royal Ontario Museum.

Description:

Morphology:

Species of Hazelia have a large variation in morphology with wide cup-shaped forms (H. palmata, H. crateria, H. luteria), long cone-shaped forms (H. conferta, H. grandis, H. obscura), branched forms (H. delicatula, H. dignata), and nodular to lobate forms (H. lobata, H. nodulifera). While there is this significant variety of overall shapes, the different species of Hazelia have a common microstructure. The walls are thin and composed of small tightly packed simple spicules that form a net-like structure and diverge outwards producing a plumose pattern. The walls are perforated with small canals to allow water flow. The base of each sponge would have had a small attachment structure.

In addition to its open shield-like shape, H. palmata possesses distinct radial tracts of spicules which go beyond the margins of the sponge for at least a couple of millimeters.

Abundance:

Hazelia is very common in the Walcott Quarry and represents 9.5% of the community (Caron and Jackson, 2008).

Maximum Size:
150 mm

Ecology:

Life habits: Epibenthic
Feeding strategies: Epibenthic
Ecological Interpretations:

Hazelia 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. 1987. Early Cambrian sponges from Vermont and Pennsylvania, the only ones described from North America. Journal of Paleontology, 61: 451-461.

RIGBY, J. K. L. F. GUNTHER AND F. GUNTHER. 1997. The first occurrence of the Burgess Shale Demosponge Hazelia palmata Walcott, 1920, in the Cambrian of Utah. Journal of Paleontology, 71: 994-997.

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

Haplophrentis carinatus

3D animation of Haplophrentis carinatus.

Animation by Phlesch Bubble © Royal Ontario Museum

Taxonomy:

Kingdom: Epibenthic
Phylum: Epibenthic
Higher Taxonomic assignment: 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:

Age:
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: Epibenthic
Feeding strategies: Epibenthic
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

Hanburia gloriosa

Hanburia gloriosa (ROM 48468). Complete individual (external mold). Specimen length = 26 mm Specimen coated with ammonium chloride sublimate to show details. Trilobite Beds on Mount Stephen.

© Royal Ontario Museum. Photo: Jean-Bernard Caron

Taxonomy:

Kingdom: Epibenthic
Phylum: Epibenthic
Higher Taxonomic assignment: Trilobita (Order: Corynexochida?)
Species name: Hanburia gloriosa
Remarks:

Trilobites are extinct euarthropods, probably stem lineage representatives of the Mandibulata, which includes crustaceans, myriapods, and hexapods (Scholtz and Edgecombe, 2006).

Described by: Walcott
Description date: 1916
Etymology:

Hanburia – unspecified, but probably after Hanbury Peak or Hanbury Glacier in the Canadian Rockies, in turn named for David T. Hanbury (1864-1910), a British explorer of the Canadian Northwest Territories.

gloriosa – from the Latin gloriosus, meaning “glorious” or “boastful,” perhaps in allusion to the unusual cephalic morphology of this rare species.

Type Specimens: Lectotype –USNM61724, in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.
Other species:

Burgess Shale and vicinity: none.

Other deposits: none.

Age & Localities:

Age:
Middle Cambrian, Glossopleura to Bathyuriscus-Elrathina Zones (approximately 505 million years ago).
Principal localities:

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

History of Research:

Brief history of research:

Walcott’s three original specimens of Hanburia gloriosa were found over the course of five years of quarrying the Phyllopod Bed on Fossil Ridge (Walcott, 1916); two more from this locality are also in theUSNMcollections. A singleUSNMspecimen was later found by Charles Resser, supposedly from the “Ogygopsis shale” on Mount Stephen (Rasetti, 1951), but this is almost certainly an error. Harry Whittington reassessed this odd trilobite in 1998.

Description:

Morphology:

Hard parts: the few known specimens of Hanburia gloriosa range in length from 4 mm (for a juvenile stage) to 35 mm. Dorsal shields are broadly ovate to subcircular in outline and all specimens are considerably flattened by compression of the thin exoskeleton. The cephalon is semicircular with a weak, shallow border furrow along the posterior and lateral margins, fading out towards the anterior corners of the glabella. The glabella in small specimens expands forwards and shows two pairs of faint bulbous lateral lobes; in larger specimens, the glabella is parallel-sided and the lobes are subdued. There are no apparent eyes located laterally on the cephalon, and there is no sign of dorsal facial suture. In these two features, Hanburia is unique among the non-agnostoid trilobites of the Burgess Shale.

Whittington (1998) has suggested that the facial suture might run along the outside edge of the cephalon, or ventrally, crossing to the dorsal side only at the genal angles, which in all specimens appear to be rounded. Larger individuals show six or seven segments in the comparatively short thorax, and a single known (presumed) juvenile stage shows four; the distal tips of the pleurae are rounded. The semicircular pygidium lacks a defined border, and is approximately the same width and length as the cephalon. Seven or eight axial rings and a terminal piece make up the pygidial axis, which ends short of the posterior margin. Eight or nine pairs of well-marked pygidial pleurae radiate out and back from the axis.

Unmineralized anatomy: not known

Abundance:

Very rare in all the Burgess Shale localities.

Maximum Size:
35 mm

Ecology:

Life habits: Epibenthic
Feeding strategies: Epibenthic
Ecological Interpretations:

Due to its unusual cephalic morphology (i.e., no dorsal sutures or lateral compound eyes), rarity, and unique occurrence only in the Burgess Shale, Hanburia gloriosa remains an ecological enigma. Other “blind” Cambrian trilobites with somewhat similar morphologies have been interpreted as inhabiting deeper waters, perhaps below the photic zone (Whittington, 1998).

References:

RASETTI, F. 1951. Middle Cambrian stratigraphy and faunas of the Canadian Rocky Mountains. Smithsonian Miscellaneous Collections, 116 (5): 1-277.

SCHOLTZ, G. AND G. D. EDGECOMBE. 2006. The evolution of arthropod heads: reconciling morphological, developmental and palaeontological evidence. Development Genes and Evolution, 216: 395-415.

WALCOTT, C. D. 1916. Smithsonian Miscellaneous Collections, 64(3): 157-258.

WHITTINGTON, H. B. 1998. Hanburia gloriosa: rare trilobite from the Middle Cambrian, Stephen Formation, British Columbia, Canada. Journal of Paleontology, 72: 673-677.

Other Links:

None

Hamptoniella foliata

Hamptoniella foliata (ROM 43816) – Holotype. Complete specimen. Specimen height = 20 mm. Specimen dry – direct light (left), wet – polarized light (right). Trilobite Beds on Mount Stephen.

© Royal Ontario Museum. Photos: Jean-Bernard Caron

Taxonomy:

Kingdom: Epibenthic
Phylum: Epibenthic
Higher Taxonomic assignment: Demospongea (Order: Monaxonida)
Species name: Hamptoniella foliata
Remarks:

Hamptoniella is a primitive demosponge, with a type of skeleton considered transitional between Hamptonia and Hazelia (Rigby, 1986). Demosponges, the same group that are harvested as bath sponges, represent the largest class of sponges today.

Described by: Rigby
Description date: 2004
Etymology:

Hamptoniella – unspecified; possibly from the town of Hampton in Virginia (see Hamptonia). The Latin suffix, ella is added to Hampton to form a diminutive.

foliata – from the Latin folia, “leaf,” in reference to the leaf like aspect of the sponge.

Type Specimens: Holotypes –ROM43816 (H. foliata wrongly referencedROM48816 in Rigby and Collins, 2004) andROM44285 (H. hirsuta) in the Royal Ontario Museum, Toronto, Canada.
Other species:

Burgess Shale and vicinity: H. hirsuta Rigby and Collins, 2004 from the Trilobite Beds on Mount Stephen.

Other deposits: none.

Age & Localities:

Age:
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:

This sponge was described by Rigby and Collins (2004) based on new material collected by the Royal Ontario Museum.

Description:

Morphology:

Hamptoniella foliata is a relatively small sponge, with its body shape varying from funnel-shaped to turbinate. The skeleton is composed of simple and straight spicules that are pointed at both ends (oxeas). The axial zone of the sponge does not have a central cavity (spongocoel) and there is no large opening (osculum). Instead a number of subvertical and relatively large canals are present in the axial area. Smaller sized canals diverge from the larger canals towards the sides and the top of the sponge. Spicules tend to be clustered and parallels to canals. H. hirsuta differs from H. foliata, by appearing more spinose.

Abundance:

H. foliata has been described based on 3 specimens and H. hirsuta based on a single specimen.

Maximum Size:
24 mm

Ecology:

Life habits: Epibenthic
Feeding strategies: Epibenthic
Ecological Interpretations:

Hamptoniella 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

Hamptonia bowerbanki

Hamptonia bowerbanki (ROM 53547). Overall view and close up of a large specimen showing the long and coarse oxeas (spicules). Specimen length = 184 mm. Specimen wet – polarized light (both images). Tulip Beds (S7) on Mount Stephen.

© Royal Ontario Museum. Photos: Jean-Bernard Caron

Taxonomy:

Kingdom: Epibenthic
Phylum: Epibenthic
Higher Taxonomic assignment: Demospongea (Order: Monaxonida)
Species name: Hamptonia bowerbanki
Remarks:

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

Described by: Walcott
Description date: 1920
Etymology:

Hamptonia – unspecified, but it comes possibly from the town of Hampton in Virginia. This town is home of the Langley Memorial Aeronautical Laboratory, which Walcott helped to create when he became first chairman of the NACA Executive Committee in 1915 (predecessor of NASA).

bowerbanki – for British naturalist and palaeontologist James Scott Bowerbank (1797-1877), best known for his studies of British sponges.

Type Specimens: Lectotype –USNM66493 (H. bowerbanki) in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA. Holotype –ROM44270 (H. elongata) in the Royal Ontario Museum, Toronto, Canada.
Other species:

Burgess Shale and vicinity: H. elongata Rigby and Collins, 2004 from the east side of Mount Field in Yoho National Park.

Other deposits: H. parva, from the Middle Cambrian Wheeler and Marjum Formations in Utah (Rigby et al., 2010); H. christi from the Lower Ordovician of Morocco (Botting, 2007).

Age & Localities:

Age:
Middle Cambrian, Bathyuriscus-Elrathina Zone to late Middle Cambrian Bolaspidella Assemblage Zone (approximately 505 million years ago).
Principal localities:

Burgess Shale and vicinity: The Walcott Quarry on Fossil Ridge. The Trilobite Beds and Tulip Beds (S7) on Mount Stephen.

Other deposits: H. bowerbanki from the Middle Cambrian Wheeler and Marjum Formations in Utah (Rigby et al., 2010).

History of Research:

Brief history of research:

Hamptonia was described by Walcott in his 1920 monograph on the sponges from the Burgess Shale. Rigby (1986) redescribed the genus, considering it to be closely related to Leptomitus and included it among the monaxial demosponges. Rigby and Collins (2004) described a new species, H. elongata, from material recently collected by the Royal Ontario Museum on Mount Field.

Description:

Morphology:

Hamptonia is a medium to large sub-hemispherical to globose sponge. The skeleton is composed of simple spicules of two sizes. Bundles or singly spaced long (up to 1 cm) coarse spicules are orientated vertically upwards away from the wall. The space between these large spicules is filled by bundle of small thatched spicules. There is a narrow central cavity and the oscular opening is circular. Faint canals are visible parallel to the long spicules that would have allowed water through the skeleton. Hamptonia may be confused with the central disc of Choia. However, Hamptonia has spicules that are much finer than Choia. H. elongata mainly differs from H. bowerbanki in that it has a branched skeleton.

Abundance:

Hamptonia bowerbanki represents only 0.09 % of the Walcott Quarry community (Caron and Jackson, 2008). Hamptonia elongata is known from a single specimen.

Maximum Size:
210 mm

Ecology:

Life habits: Epibenthic
Feeding strategies: Epibenthic
Ecological Interpretations:

Hamptonia 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:

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.

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

Hallucigenia sparsa

3D animation of Hallucigenia sparsa.

Animation by Phlesch Bubble © Royal Ontario Museum

Taxonomy:

Kingdom: Epibenthic
Phylum: Epibenthic
Higher Taxonomic assignment: Xenusia (Order: Scleronychophora, stem group onychophorans)
Species name: Hallucigenia sparsa
Remarks:

Hallucigenia is regarded as a member of the “lobopodans,” a group of vermiform Cambrian organisms possessing pairs of leg-like extensions of the body. The affinities of these animals are controversial; they have been placed at the base of a clade comprised of anomalocaridids and arthropods (Budd, 1996), or in a stem-group to modern onychophorans (Ramsköld and Chen, 1998).

Described by: Walcott
Description date: 1911
Etymology:

Hallucigenia – from the Latin hallucinatio, “wandering of the mind,” after the bizarreness of the animal.

sparsa – from the Latin sparsus, “rare, or scattered,” reflecting the rarity of the specimens available in the original study.

Type Specimens: Holotype –USNM83935 in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.
Other species:

Burgess Shale and vicinity: none.

Other deposits: H. fortis from the Middle Cambrian Chengjiang biota (Hou and Bergström 1995).

Age & Localities:

Age:
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:

Hallucigenia was originally described as “Canadia sparsa” by Walcott (1911) in a review of various Burgess Shale “annelids.” One specimen was illustrated twenty years later (Walcott, 1931), but the first thorough study of this animal wasn’t published until Conway Morris (1977) demonstrated that it did not belong to the genus Canadia or to the annelids at all. His reconstruction showed a bizarre animal walking on spines, with dorsal tentacles interpreted as a feeding apparatus (Conway Morris, 1977). The new genus name Hallucigenia was coined in reference to this “dreamlike” appearance and also reflected the organism’s uncertain affinities. It was later shown that the supposed tentacles represented just one row of paired “legs” – the others were buried under a layer of rock and the paired spines were on the dorsal surface (Ramsköld and Hou, 1991, Ramsköld, 1992). The anteroposterior orientation was also reversed, with the former head interpreted as possible decay fluids seeping from the body (Ramsköld, 1992).

Description:

Morphology:

Hallucigenia has a worm-like body with a small head at the end of a long neck; the trunk bears seven pairs of long dorsal spines and seven pairs of slender leg-like lobes. The spacing between lobes and spines is relatively constant. The spine pairs are shifted forward so that the posterior pair of legs does not have a corresponding pair of spines above. Each leg terminates in a pair of claws and the rigid spines have inflexible basal plates. The neck area bears two or three pairs of very fine anterior “appendages” lacking terminal claws. The head is indistinct but the mouth is anterior; a straight gut ends in a posterior anus. It is possible the posterior end is in fact more bulbous than previously thought.

Abundance:

About thirty specimens were studied by Conway Morris (1977). Overall, Hallucigenia is rare, and in the Walcott Quarry it represents 0.19% of the specimens counted in the community (Caron and Jackson, 2008).

Maximum Size:
30 mm

Ecology:

Life habits: Epibenthic
Feeding strategies: Epibenthic
Ecological Interpretations:

Hallucigenia is often found in association with the sponge Vauxia and other organic debris. This co-occurrence has led to suggestions that Hallucigenia fed on sponges, using its clawed legs to hang on, with its spines protecting it from predation. It is also possible that Hallucigenia scavenged on decaying animal remains.

References:

BUDD, G. E. 1996. The morphology of Opabinia regalis and the reconstruction of the arthropod stem-group. Lethaia, 29: 1-14.

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. 1977. A new metazoan from the Burgess Shale of British Columbia. Palaeontology, 20: 623-640.

CONWAY MORRIS, S. 1999. The crucible of creation: the Burgess Shale and the rise of animals. Oxford University Press, USA.

HOU, X. AND J. A. N. BERGTRÖM. 1995. Cambrian lobopodians – ancestors of extant onychophorans? Biological Journal of the Linnean Society, 114(1): 3-19.

RAMSKÖLD, L. 1992. The second leg row of Hallucigenia discovered. Lethaia, 25(2): 221–224.

RAMSKÖLD, L. AND X. HOU. 1991. New early Cambrian animal and onychophoran affinities of enigmatic metazoans. Nature, 351: 225-228.

RAMSKÖLD, L. AND J. Y. CHEN. 1998. Cambrian lobopodians: morphology and phylogeny, p. 107-150. In G. D. Edgecombe (ed.), Arthropod fossils and phylogeny. Volume 29. Columbia University Press, New York.

WALCOTT, C. 1911. Cambrian Geology and Paleontology II. Middle Cambrian annelids. Smithsonian Miscellaneous Collections, 57(5): 109-145.

WALCOTT, C. 1931. Addenda to descriptions of Burgess Shale fossils. Smithsonian Miscellaneous Collections, 85(3): 1-46.

Other Links:

http://gsc.nrcan.gc.ca/paleochron/09_e.php

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

http://evolution.berkeley.edu/evolibrary/article/_0_0/cambrian_13

Halichondrites elissa

Halichondrites elissa (ROM 53575) – Part and counterpart. Nearly complete individual showing the pointed root tuft. Specimen height = 140 mm. Specimen dry – direct light (left), wet – direct light (right). Walcott Quarry.

© Royal Ontario Museum. Photos: Jean-Bernard Caron

Taxonomy:

Kingdom: Epibenthic
Phylum: Epibenthic
Higher Taxonomic assignment: Demospongea (Order: Monaxonida)
Species name: Halichondrites elissa
Remarks:

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

Described by: Walcott
Description date: 1920
Etymology:

Halichondrities – from the Greek hal, meaning “belonging to the sea,” chon, meaning “funnel” or “tube,” and dri, meaning “thicket.” The name refers to the shape of this marine sponge with a thicket of long hair-like spicules.

elissa – from the Greek eliss, meaning “to roll, or to turn about.” This name may refer to the spiral pattern of the small spicules of this sponge.

Type Specimens: Holotype –USNM66447 in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.
Other species:

Burgess Shale and vicinity: An unidentified species, Halichondrites sp. from Mount Stephen (Rigby and Collins, 2004).

Other deposits: H. confusus Dawson, 1889 from the Ordovician of Quebec at Little Métis.

Age & Localities:

Age:
Lower Cambrian, Wutingaspis-Eoredlichia Zone to Middle Cambrian, Bathyuriscus-Elrathina Zone (approximately 510 to 505 million years ago).
Principal localities:

Burgess Shale and vicinity: The Walcott and Raymond Quarries on Fossil Ridge.

Other deposits: H. elissa from the Lower Cambrian Chengjiang fauna (Chen et al., 1997; Luo et al., 1999).

History of Research:

Brief history of research:

Walcott assigned this species to the genus Halichondrites in 1920. Ribgy (1986) re-described this genus and hypothesized that Halichondrites probably evolved from an early species of Leptomitus and established a new family called Halichondritidae to include this genus. New specimens collected by the Royal Ontario Museum were subsequently described by Rigby and Collins in 2004.

Description:

Morphology:

This sponge has a cone shaped base that extends upwards to form a long tube. The walls of the sponge are smooth with a thatch of small spicules that are vertically arranged in a clockwise spiraling pattern. There are no canals visible in the wall; they may be very small or run parallel to the wall. The most distinctive part of this sponge is the long thick, densely arranged spicules that emerge from the wall. These spicules are orientated upwards and may be up to 8.5 cm long. This sponge can be over 20 cm tall and is one of the tallest and most hirsute (densely covered in hair) of the Burgess Shale sponges. Water would have entered though small pores in the wall, moving into the central cavity and out the circular osculum at the top of the sponge.

Abundance:

Halichondrites is very rare and represents only 0.01% of the Walcott Quarry community (Caron and Jackson, 2008).

Maximum Size:
215 mm

Ecology:

Life habits: Epibenthic
Feeding strategies: Epibenthic
Ecological Interpretations:

Halichondrites 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.

CHEN, J. Y., Y. N. CHENG AND H. V. ITEN. 1997. The Cambrian explosion and the fossil record. National Museum of Natural Science Taiwan, Taichung, 319 p.

LUO, H., S. HU, L. CHEN, S. ZHANG AND Y. TAO. 1999. Early Cambrian Chengjiang fauna from Kunming region, China. Yunnan Science and Technology Press, Kunming, 162 p.

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