The Burgess Shale

Hurdia victoria

3D animation of Hurdia victoria.

Animation by Phlesch Bubble © Royal Ontario Museum

Taxonomy:

Kingdom: Others
Phylum: Others
Class: Dinocarida (Order: Radiodonta, stem group arthropods)
Species name: Hurdia victoria
Remarks:

Hurdia is an anomalocaridid, and is usually considered to represent either a basal stem-lineage euarthropod (e.g. Daley et al., 2009), a member of the crown-group arthropods (e.g. Chen et al., 2004), or a sister group to the arthropods (Hou et al., 2006).

Described by: Walcott
Description date: 1912
Etymology:

Hurdia – from Mount Hurd (2,993 m), a mountain northeast of the now defunct Leanchoil railway station on the Canadian Pacific Railway in Yoho National Park. The peak was named by Tom Wilson for Major M. F. Hurd, a CPR survey engineer who explored the Rocky Mountain passes starting in the 1870s.

victoria – unspecified; perhaps from Mount Victoria (3,464 m) on the border of Yoho and Banff National Parks, named by Norman Collie in 1897 to honour Queen Victoria.

Type Specimens: Lectotypes –USNM57718 (H. victoria) andUSNM57721 (H. triangulata) in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.
Other species:

Burgess Shale and vicinity: Hurdia triangulata.

Other deposits: Potentially other species are represented in Utah (Wheeler Formation) (Briggs et al., 2008), the Jince Formation in the Czech Republic (Chlupáč and Kordule 2002) and the Shuijingtuo Formation in Hubei Province, China (Cui and Huo, 1990) and possibly Nevada (Lieberman, 2003).

Age & Localities:

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

The Walcott, Raymond and Collins Quarries on Fossil Ridge. Also known from other localities on Mount Field, Mount Stephen – Tulip Beds (S7) – and near Stanley Glacier.

History of Research:

Brief history of research:

Hurdia is a relative newcomer to the anomalocaridids. Although isolated parts of its body were first identified in the early 1900s, no affinity could be determined until the description of whole body specimens by Daley et al. in 2009. Hurdia victoria was the name originally given to an isolated triangular carapace that Walcott (1912) suggested belonged to an unknown arthropod. Proboscicaris, another isolated carapace, was originally described as a phyllopod arthropod (Rolfe, 1962). Hurdia’s frontal appendages were first described by Walcott (1911a) as the feeding limbs of Sidneyia, but were later removed from this genus and referred to as “Appendage F” with unknown affinity (Briggs, 1979).

Like other anomalocaridids, the mouth parts were first described as the jellyfish Peytoia nathorsti (Walcott, 1911b). When Whittington and Briggs (1985) discovered the first whole body specimens of Anomalocaris, the mouth part identity of Peytoia was recognized and “Appendage F” was determined to be the frontal appendage of Anomalocaris nathorsti (later renamed Laggania cambria by Collins (1996). When describing Anomalocaris, Whittington and Briggs (1985) also figured a mouth apparatus with extra rows of teeth.

After two decades of collecting at the Burgess Shale, Desmond Collins from the Royal Ontario Museum (ROM) discovered that this extra-spiny mouth part actually belonged to a third type of anomalocaridid, which also had an “Appendage F” pair and a frontal carapace structure consisting of one Hurdia carapace and two Proboscicaris carapaces (Daley et al., 2009). This is the Hurdia animal. ROM specimens of “Appendage F” showed that it has three distinct morphologies, two of which belongs to the Hurdia animal (known from two species, victoria and triangulata) and one to Laggania cambria.

Description:

Morphology:

Hurdia has a bilaterally symmetrical body that is broadly divisible into two sections of equal lengths. The anterior region is a complex of non-mineralized carapaces consisting of one dorsal triangular H-element (previously called Hurdia) and two lateral subrectangular P-elements (or Proboscicaris). This complex is hollow and open ventrally. It attaches near the anterior margin of the head and protrudes forward. The surfaces of the H- and P-elements are covered in a distinctive polygonal pattern similar to that seen on Tuzoia carapaces. A pair of oval eyes on short stalks protrudes upwards through dorsal-lateral notches in the overlapping posterior corners of the H- and P-elements.

Mouth parts are on the ventral surface of the head, and consist of a circlet of 32 tapering and overlapping plates, 4 large and 28 small, with spines lining the square inner opening. Within the central opening are up to five inner rows of toothed plates. A pair of appendages flanks the mouth part, each with nine thin segments with short dorsal spines and seven elongated ventral spines. The posterior half of the body consists of a series of seven to nine reversely imbricated lateral lobes that extend ventrally into triangular flaps. Dorsal surfaces of the lateral lobes are covered in a series of elongated blades interpreted to be gill structures. The body terminates abruptly in two rounded lobes, and lacks a tailfan. Complete specimens are up to 20 cm in length, although disarticulated fragments may suggest a larger body size up to 50 cm long. Hurdia triangulata differs from Hurdia victoria by having a wider and shorter H-element.

Abundance:

Over 700 specimens of Hurdia have been identified, most of which are disarticulated. Hurdia is found in all Burgess Shale quarries on Fossil Ridge, and is particularly abundant in Raymond Quarry, where it makes up almost 1% of the community (240 specimens). A total of 7 complete body specimens exist.

Maximum Size:
500 mm

Ecology:

Life habits: Others
Feeding strategies: Others
Ecological Interpretations:

Hurdia was likely nektonic, since there are no trunk limbs for walking, and the numerous gills suggest an active swimming lifestyle. The animal propelled itself through the water column by waving its lateral lobes and gills. The large eyes, prominent appendages and spiny mouth parts suggest that Hurdia actively sought out moving prey items. Although the function of the frontal carapace remains unknown, it may have played a role in prey capture. If Hurdia were swimming just above the sea floor, it could have used the tip of its frontal carapace to stir up sediment and dislodge prey items, which would then be trapped beneath its frontal carapace. Prey items were funneled towards the mouth by a sweeping motion of the long ventral blades of the frontal appendages, which formed a rigid net or cage. Like other anomalocaridids, Hurdia likely ingested soft-bodied prey.

References:

BRIGGS, D. E. G. 1979. Anomalocaris, the largest known Cambrian arthropod. Palaeontology, 22: 631-663.

BRIGGS, D. E. G., B. S. LIEBERMAN, J. R. HENDRICK, S. L. HALGEDAHL AND R. D. JARRARD. 2008. Middle Cambrian arthropods from Utah. Journal of Paleontology, 82: 238-254.

CHEN, J. Y., D. WALOSZEK AND A. MAAS. 2004. A new ‘great-appendage’ arthropod from the Lower Cambrian of China and homology of chelicerate chelicerae and raptorial antero-ventral appendages. Lethaia, 37: 3-20.

CHLUPÁČ, I. AND V. KORDULE. 2002. Arthropods of Burgess Shale type from the Middle Cambrian of Bohemia (Czech Republic). Bulletin of the Czech Geological Survey, 77: 167-182.

COLLINS, D. 1996. The “evolution” of Anomalocaris and its classification in the arthropod class Dinocarida (nov) and order Radiodonta (nov). Journal of Paleontology, 70: 280-293.

CUI, Z. AND S. HUO. 1990. New discoveries of Lower Cambrian crustacean fossils from Western Hubei. Acta Palaeontologica Sinica, 29: 321-330.

DALEY, A. C., G. E. BUDD, J. B. CARON, G. D. EDGECOMBE AND D. COLLINS. 2009. The Burgess Shale anomalocaridid Hurdia and its significance for early euarthropod evolution. Science, 323: 1597-1600.

HOU, X., J. BERGSTRÖM AND P. AHLBERG. 1995. Anomalocaris and other large animals in the Lower Cambrian Chengjiang fauna of Southwest China. GFF, 117: 163-183.

HOU, X., J. BERGSTRÖM AND Y. JIE. 2006. Distinguishing anomalocaridids from arthropods and priapulids. Geological Journal, 41: 259-269.

LIEBERMAN, B. S. 2003. A new soft-bodied fauna: The Pioche Formation of Nevada. Journal of Paleontology, 77: 674-690.

ROLFE, W. D. I. 1962. Two new arthropod carapaces from the Burgess Shale (Middle Cambrian) of Canada. Breviora Museum of Comparative Zoology, 60: 1-9.

WALCOTT, C. D. 1911a. Middle Cambrian Merostomata. Cambrian Geology and Paleontology II. Smithsonian Miscellaneous Collections, 57: 17-40.

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

WALCOTT, C. D. 1912. Middle Cambrian Branchiopoda, Malacostraca, Trilobita and Merostomata. Smithsonian Miscellaneous Collections, 57: 145-228.

WHITTINGTON, H. B. AND D. E. G. BRIGGS. 1985. The largest Cambrian animal, Anomalocaris, Burgess Shale, British-Columbia. Philosophical Transactions of the Royal Society of London Series B-Biological Sciences, 309: 569-609.

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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: Others
Phylum: Others
Class: 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:

Period:
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: Others
Feeding strategies: Others
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: Others
Phylum: Others
Class: Hyolitha (Order: Hyolithida, stem group molluscs)
Species name: Haplophrentis carinatus
Remarks:

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

Described by: Matthew
Description date: 1899
Etymology:

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

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

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

Burgess Shale and vicinity: none

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

Age & Localities:

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

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

History of Research:

Brief history of research:

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

Description:

Morphology:

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

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

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

Abundance:

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

Maximum Size:
40 mm

Ecology:

Life habits: Others
Feeding strategies: Others
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.

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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: Others
Phylum: Others
Class: 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:

Period:
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: Others
Feeding strategies: Others
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

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: Others
Phylum: Others
Class: 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:

Period:
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: Others
Feeding strategies: Others
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:

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Pollingeria grandis

Pollingeria grandis (GSC 8362). Slab with several specimens. Specimen length (largest) = 15 mm. Specimen dry – polarized light. Walcott Quarry.

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

Taxonomy:

Kingdom: Others
Phylum: Others
Class: Non applicable
Species name: Pollingeria grandis
Remarks:

Pollingeria is one of the least understood Burgess Shale organisms, and its systematic status is unknown (Briggs and Conway Morris, 1986).

Described by: Walcott
Description date: 1911
Etymology:

Pollingeria – from Mount Pollinger (2,816 m), northwest of the Burgess Shale. The name was given after Joseph Pollinger (1873-1943).

grandis – from the Latin grandis, “big, large,” in reference to the purported large size of the fossils.

Type Specimens: Syntypes –USNM57639-57641 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 and smaller sites on Mount Field and Mount Stephen.

History of Research:

Brief history of research:

Pollingeria was first described by Walcott in a 1911 monograph dealing with various Burgess Shale worms. Walcott interpreted these fossils as the individual scales of a larger organism resembling Wiwaxia. However, this interpretation was doubted (Conway Morris, 1979), and firmly rejected after the restudy of Wiwaxia (Conway Morris, 1985). The affinities of Pollingeria have remained difficult to establish (Briggs and Conway Morris, 1986).

Description:

Morphology:

The shape of this fossil is ovoid but variable in details and most individuals range from 10 to 15 mm in length. A distinctive feature is a series of narrow tubular elements that are darker and often slightly raised; these are twisted and contorted and do not appear to be parts of a gut.

Abundance:

Pollingeria is locally very abundant with hundreds of specimens on some bedding surfaces. In the Walcott Quarry this species represents 5.83% of the specimens counted in the community (Caron and Jackson, 2008).

Maximum Size:
15 mm

Ecology:

Life habits: Others
Feeding strategies: Others
Ecological Interpretations:

Not enough is known about this organism to interpret its ecology.

References:

BRIGGS, D. E. G. AND S. CONWAY MORRIS. 1986. Problematica from the Middle Cambrian Burgess Shale of British Columbia, p. 167-183. In A. Hoffman and M. H. Nitecki (eds.), Problematic fossil taxa (Oxford Monographs on Geology and Geophysics No. 5). Oxford University Press & Clarendon Press, New York.

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

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

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

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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:

Kingdom: Others
Phylum: Others
Class: Demospongea (Order: Monaxonida)
Species name: Pirania muricata
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.

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:

Life habits: Others
Feeding strategies: Others
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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Fieldospongia bellilineata

Fieldospongia bellilineata (ROM 53602). Complete specimen showing root tuft to upper margin. Specimen height = 55 mm. Specimen wet – polarized light. The Monarch Cirque.

© Royal Ontario Museum. Photo: Jean-Bernard Caron

Taxonomy:

Kingdom: Others
Phylum: Others
Class: Demospongea (Order: Monaxonida)
Species name: Fieldospongia bellilineata
Remarks:

Fieldospongia is considered to be a primitive demosponge related to the Anthaspidellidae (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:

Fieldospongia – from Field, the mountain peak (2,643 m) and small town near Fossil Ridge, British Columbia, Canada. The name was given by William Cornelius Van Horne (General Manager of the Canadian Pacific Railway), to honor Cyrus West Field a promoter of the first telegraph cable across the Atlantic Ocean; and from the Latin spongia, “sponge.”

bellilineata – from the Latin bell, “charming,” and linea, “pertaining to lines.” The name makes reference to the linear-like skeleton of this sponge.

Type Specimens: Holotype –USNM66454, 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, probably from the Glossopleura Zone or the Bathyuriscus-Elrathina Zone (approximately 505 million years ago).
Principal localities:

The northwest shoulder of Mount Stephen and the Monarch Peak in Kootenay National Park.

History of Research:

Brief history of research:

Described in 1920 as Tuponia bellilineata by Walcott in his monograph on sponges from the Burgess Shale, this form has now been placed within a separate genus, Fieldospongia, erected by Rigby in 1986. The only known specimen purportedly came from the Mount Whyte Formation (Walcott, 1920), just below the Northwestern shoulder of Mount Stephen. This formation is older than any of the rock units containing Burgess Shale-type fossils in the vicinity and is not characterized by exceptional preservation. Examination of the specimen suggests that this sponge probably comes from younger rock units with Burgess Shale-type fossils (Glossopleura to Bathyuriscus-Elrathina Zones) that are known in this area. It is likely that this specimen was derived from upper sections and was collected at the base of talus slopes coinciding with levels of the Mount Whyte Formation. A second specimen was described by Rigby and Collins in 2004 from The Monarch, a mountain Peak at the border between Mount Assiniboine Provincial Park and Kootenay National Park.

Description:

Morphology:

This species has a conical shape with gentle horizontal wrinkles. The thin walls are composed of similarly spaced bundles of spicules arranged vertically and horizontally, forming a regular skeletal net with rectangular cells. Vertical strands are usually more prominent than the horizontal ladders. The root tuft is relatively long and narrow.

Abundance:

Fieldospongia is very rare, known from only two specimens.

Maximum Size:
55 mm

Ecology:

Life habits: Others
Feeding strategies: Others
Ecological Interpretations:

Fieldospongia 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:

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Elrathina cordillerae

Elrathina cordillerae (ROM 53273). Complete individual; a presumed carcass with free cheeks in place (coated with ammonium chloride sublimate to show details). Specimen length = 24 mm. Specimen dry – direct light. Mount Stephen Trilobite Beds on Mount Stephen.

© Royal Ontario Museum. Photo: Jean-Bernard Caron

Taxonomy:

Kingdom: Others
Phylum: Others
Class: Trilobita (Order: Ptychopariida)
Species name: Elrathina cordillerae
Remarks:

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

Described by: Rominger
Description date: 1887
Etymology:

Elrathina – unspecified.

cordillerae – in reference to the Western Cordillera (Canadian Rocky Mountain ranges), derived from the Spanish cordilla, the diminutive of cuerda, meaning “cord.”

Type Specimens: Type status under review – UMMP 4883 (6 specimens), University of Michigan Museum of Paleontology, Ann Arbor, Michigan, USA.
Other species:

Burgess Shale and vicinity: Elrathina parallela, E. brevifrons, E. spinifera, and E. marginalis have been described from similar stratigraphic horizons at nearby sites on Mount Field, Mount Stephen, and Mount Odaray.

Other deposits: Other species of Elrathina have been reported from the Cambrian of North America and Greenland.

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 additional localities on Mount Stephen.

History of Research:

Brief history of research:

E. cordillerae was originally described under the genus name Conocephalites in Rominger’s 1887 publication on trilobites from Mount Stephen. In 1888 Walcott reallocated the species to Ptychoparia where it remained until Charles Resser, Walcott’s former assistant at the United States National Museum, established the new replacement genus Elrathina (Resser, 1937). Other workers have subsequently suggested that Elrathina is indistinguishable from Ptychoparella (see Blaker and Peel, 1997).

Species of Elrathina, along with those of the corynexochid Bathyuriscus, were found to be very abundant in a narrow interval of Middle Cambrian rocks throughout western North America, forming the basis of the Bathyuriscus-Elrathina Zone erected by Charles Deiss (1940).

Description:

Morphology:

Hard parts: adult dorsal exoskeletons average about 2 cm long. The semicircular cephalon is about one-third the length of the entire dorsal shield, bordered by a well-defined narrow rim, and with rounded genal angles. Weak transverse eye ridges extend to the small eyes, which are located just forward of cephalic mid-length. The slightly anteriorly narrowing glabella is rounded in front and exhibits three pairs of shallow lateral furrows; the pre-glabellar field is about the same width as the narrow anterior rim. The long, tapering thorax with a narrow axial lobe contains between 17 and 19 straight-sided segments, flexed gently downwards a short distance from the rounded tips. The tiny elliptical pygidium usually features two segments.

Unmineralized anatomy: rare specimens from the Walcott Quarry on Fossil Ridge retain tantalizing evidence of soft parts, including a pair of slender uniramous antennae, followed by very delicate looking biramous limbs beneath the cephalon, thorax and pygidium. These and other individuals of E. cordillerae are occasionally associated with a dark stain adjacent to the exoskeleton, presumably representing fluidized decay products.

Abundance:

Relatively common on Fossil Ridge and locally very abundant in the Walcott Quarry, where it represents about 25% of all trilobites collected (Caron and Jackson, 2008).

Maximum Size:
28 mm

Ecology:

Life habits: Others
Feeding strategies: Others
Ecological Interpretations:

Like similar-looking ptychoparioid trilobites, E. cordillerae may be interpreted as a fully mobile, epibenthic deposit (particle) feeder adapted to very low oxygen levels.

References:

BLAKER, M. R. AND J. S. PEEL. 1997. Lower Cambrian trilobites from North Greenland. Meddeleser om Grønland, Geoscience, 35, 145 p.

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

DEISS, C. 1940. Lower and Middle Cambrian stratigraphy of southwestern Alberta and southeastern British Columbia. Bulletin of the Geological Society of America, 51: 731-794.

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

RESSER, C. E. 1937. Third contribution to nomenclature of Cambrian trilobites. Smithsonian Miscellaneous Collections, 95(22): 29 p.

ROMINGER, C. 1887. Description of primordial fossils from Mount Stephens, N. W. Territory of Canada. Proceedings of the Academy of Natural Sciences of Philadelphia, 1887: 12-19.

RUDKIN, D. M. 1989. Trilobites with appendages from the Middle Cambrian Stephen Formation of British Columbia. 28th International Geological Congress, Washington, D.C. July 9-19, 1989. Abstracts: 2-729.

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. 1918. Cambrian Geology and Paleontology IV. Appendages of trilobites. Smithsonian Miscellaneous Collections, 67(4): 115-216.

WALCOTT, C. D. 1924. Cambrian and Lower Ozarkian trilobites. Smithsonian Miscellaneous Collections, 75(2): 53-60.

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Elrathia permulta

Elrathia permulta (ROM 60762). Complete individual; a presumed carcass with free cheeks in place. Specimen length = 27 mm. Specimen dry – direct light (left) and coated with ammonium chloride sublimate to show details (right). Walcott Quarry.

© Royal Ontario Museum. Photo: Jean-Bernard Caron

Taxonomy:

Kingdom: Others
Phylum: Others
Class: Trilobita (Order: Ptychopariida)
Species name: Elrathia permulta
Remarks:

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

Described by: Walcott
Description date: 1918
Etymology:

Elrathia – unspecified.

permulta – from the Latin per, “very much”, and multus, “many”.

Type Specimens: Holotype –USNM65517 in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.
Other species:

Burgess Shale and vicinity: none.

Other deposits: other species occur, sometimes abundantly, elsewhere in the Cambrian of North America and Greenland.

Age & Localities:

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

The Walcott Quarry on Fossil Ridge. Elrathia sp. has been reported from localities on Mount Stephen.

History of Research:

Brief history of research:

The concept of Elrathia permulta is quite confused. Walcott named the species Ptychoparia permulta in 1918 and illustrated two specimens, one clearly designated as the type. Resser (1937) noted that the illustrated specimens were quite different, moved both to Walcott’s 1924 genus Elrathia, and proposed the name Elrathia dubia for the second species. Unfortunately, he based this on the original type of permulta; Rasetti (1951) declared dubia invalid, returned the type specimen to Elrathia permulta, and designated the other of Walcott’s specimens as a paratype of Ehmaniella burgessensis. The holotype of permulta, however, lacks many of the diagnostic characters of Elrathia, and Robison (1964) suggested it represents a new genus.

Description:

Morphology:

Hard parts: the adult dorsal exoskeleton is up to 25 mm long, with a large semicircular cephalon occupying about one-third the total length. The cephalon is bordered by a rounded rim and broad inner furrow; genal angles are produced into sharp triangular spines extending back to the fourth thoracic segment. There is a relatively long field between the narrow, tapered, and anteriorly rounded glabella and the frontal rim. Eyes are small and transverse eye ridges are very weak. Three pairs of shallow lateral furrows mark the glabella. The thorax comprises 14 segments, and tapers back more rapidly over the posterior half to a small rounded pygidium. The surface of the exoskeleton is variably granulate.

Unmineralized anatomy: not known.

Abundance:

Rare in the Walcott Quarry on Fossil Ridge, and elsewhere.

Maximum Size:
25 mm

Ecology:

Life habits: Others
Feeding strategies: Others
Ecological Interpretations:

E. permulta may, like similar small ptychoparioid trilobites, be interpreted as a mobile, epibenthic deposit (particle) feeder adapted to low oxygen levels.

References:

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

RESSER, C. E. 1937. Third contribution to nomenclature of Cambrian trilobites. Smithsonian Miscellaneous Collections, 95(22): 29 p.

ROBISON, R. A. 1964. Late Middle Cambrian faunas from western Utah. Journal of Paleontology, 38:510-566.

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. 1918. Cambrian Geology and Paleontology IV. Appendages of trilobites. Smithsonian Miscellaneous Collections, 67(4): 115-216.

WALCOTT, C. D. 1924. Cambrian and Lower Ozarkian trilobites. Smithsonian Miscellaneous Collections, 75(2): 53-60.

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