The Burgess Shale

Surusicaris elegans

Surusicaris elegans, holotype ROMIP 62976. Specimen dry – direct light (left column), dry – polarized light (right column).

Taxonomy:

Kingdom: Extremely rare
Phylum: Extremely rare
Higher Taxonomic assignment: Family Isoxyidae?
Species name: Surusicaris elegans
Remarks:

Surusicaris is a close relative of Isoxys, as indicated by the type of carapace, eyes, and frontal pair of raptorial appendages (Aria & Caron, 2015). The presence of spines on the dorsal side of the frontal appendage is a character shared with radiodontans, such as Anomalocaris. Current evidence draws out a consensus among authors placing isoxyids as sister taxa to true arthropods (Edgecombe, 2020; Aria, 2022), although it is not clear whether Surusicaris and Isoxys are part of a single separate lineage (that is, form a monophyletic group).

Described by: Aria and Caron
Description date: 2015
Etymology:

Surusicaris – After Surus, “the Syrian,” which would have been the last elephant of Hannibal, with broad shields covering its sides and missing a tusk.

elegans – Referring to the delicate, laced appearance of the limbs.

Type Specimens: Holotype ROMIP 62977, at the Royal Ontario Museum, Toronto, Canada
Other species:

Burgess Shale and vicinity: None
Other deposits: None

Age & Localities:

Age:
Middle Cambrian, Wuliuan stage, upper part of the Burgess Shale Formation (around 507 million years old).
Principal localities:

Marble Canyon, Kootenay National Park, British Columbia.

History of Research:

Brief history of research:

Along with Yawunik kootenayi (Aria, Caron & Gaines, 2015), Surusicaris elegans (Aria & Caron, 2015) is one of the first two new arthropods described from the Marble Canyon locality of the Burgess Shale. The original study was based on a single specimen from the original 2012 expedition. No other specimen has been confirmed so far, in the Burgess Shale or elsewhere. Surusicaris has remained a critical taxon in understanding the place of isoxyids in the transition to a euarthropod body plan (Fu et al., 2022; Aria, 2022).

Description:

Morphology:

Surusicaris elegans is about 15mm long and enclosed in a broad carapace made of two semi-circular valves, without spines. Only the posterior extremity of the body and tailpiece remain uncovered. The animal has a well-defined head composed of, at the front, a pair of large spherical eyes and a segmented predatory appendage, and, at the back of the head, under the carapace, three pairs of short limbs with a lobopod aspect. The frontal appendages show a complex ornament of spines on both the ventral and dorsal margins. The trunk limbs are clearly bipartite, forming two separate but similar branches. As for Isoxys, external segmentation of the trunk is not clearly visible. Inside the body, a bold, black trace runs alongside the gut and branches out inside one limb branch, showing similarities to hemolymphatic (“blood”) channels (Aria & Caron, 2015).

Abundance:

A single specimen from the Marble Canyon quarry.

Maximum Size:
About 15 mm.

Ecology:

Life habits: Extremely rare
Feeding strategies: Extremely rare
Ecological Interpretations:

The large lateral eyes and clawed frontal appendages suggest Surusicaris was an active predator, like its close relative Isoxys (Legg & Vannier, 2013). Trunk limbs lack strong functional modifications, but their lobate aspect in addition to their position underneath the carapace indicates that Surusicaris was mostly a swimmer (Aria & Caron, 2015). Although some authors implied a pelagic lifestyle (Vannier & Chen, 2000), isoxyids are commonly found among benthic/nektobenthic assemblages (Caron & Jackson, 2008) and possess general morphological characteristics of other nektobenthic Cambrian arthropods.

References:

  • Aria, C. (2022) The origin and early evolution of arthropods. Biological Reviews 97, 1786–1809.
  • Aria, C. & Caron, J.-B. (2015) Cephalic and limb anatomy of a new isoxyid from the Burgess Shale and the role of ‘stem bivalved arthropods’ in the disparity of the frontalmost appendage. PLoS ONE 10, e0124979.
  • Aria, C., Caron, J.-B. & Gaines, R. (2015) A large new leanchoiliid from the Burgess Shale and the influence of inapplicable states on stem arthropod phylogeny. Palaeontology 58, 629–660.
  • Caron, J.B. & Jackson, D.A. (2008) Paleoecology of the Greater Phyllopod Bed community, Burgess Shale. Palaeogeography, Palaeoclimatology, Palaeoecology 258, 222–256.
  • Edgecombe, G.D. (2020) Arthropod origins: Integrating paleontological and molecular evidence. Annual Review of Ecology, Evolution, and Systematics 51, 1–25.
  • Fu, D., Legg, D.A., Daley, A.C., Budd, G.E., Wu, Y. & Zhang, X. (2022) The evolution of biramous appendages revealed by a carapace-bearing Cambrian arthropod. Philosophical Transactions of the Royal Society B: Biological Sciences 377, 20210034.
  • Legg, D.A. & Vannier, J. (2013) The affinities of the cosmopolitan arthropod Isoxys and its implications for the origin of arthropods. Lethaia 46, 540–550.
  • Vannier, J. & Chen, J.Y. (2000) The Early Cambrian colonization of pelagic niches exemplified by Isoxys (Arthropoda). Lethaia 33, 295–311.
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Collinsovermis monstruosus

Collinsovermis monstruosus, holotype ROMIP 52703

Taxonomy:

Kingdom: Extremely rare
Phylum: Extremely rare
Higher Taxonomic assignment: Order Luolishaniida, Family Collinsovermidae
Species name: Collinsovermis monstruosus
Remarks:

Collinsovermis is one of a variety of lobopodian taxa from the Cambrian, which are early members of the lineage that gave rise to arthropods, and whose only modern survivors are onychophorans (velvet worms) and tardigrades (water bears). Lobopodians characteristically have annulated, unjointed bodies and bear soft limbs after which they are called: the lobopods. Collinsovermis is an armoured member of the Order Luolishaniida, along with forms such as Collinsium and Luolishania from China, or Acinocricus from Utah—together forming the family Collinsovermidae. Luolishaniids are characterized by their thin spines arranged in chevrons and the differentiation of their body into functional regions for suspension-feeding (Caron & Aria 2017, 2020).

Described by: Caron and Aria
Description date: 2020
Etymology:

Collinsovermis – Collins, patronymic, honours its discoverer, Desmond Collins, and vermis is Latin for worm.

monstruosus – From the Latin, in reference to the nickname ‘Collins’ monster’, first introduced by Delle Cave & Simonetta (1991).

Type Specimens: Holotype: ROMIP 52703, Paratypes: ROMIP 52704 and 52705 at the Royal Ontario Museum, Toronto, Canada.
Other species:

Burgess Shale and vicinity: None.
Other deposits: None.

Age & Localities:

Age:
Middle Cambrian, Wuliuan stage, Bathyuriscus-Elrathina Zone of the Burgess Shale Formation (approximately 507 million years old).
Principal localities:

Mount Stephen

History of Research:

Brief history of research:

Collected by a Royal Ontario Museum expedition on Mount Stephen in 1983, the Collinsovermis animal was first revealed, unnamed and undescribed, in the non-peer-reviewed magazine Rotunda (former name of the Royal Ontario Museum Magazine), as a single picture of the holotype, with the caption: “What is it? This new spiny animal (4 cm) with hairy legs has a body plan that has not been seen before.” In 1991, Italian scientists Delle Cave and Simonetta (1991) provided a brief description of the taxon and attempted a reconstruction solely based on the photograph provided by Collins in the Rotunda magazine, coining it the “Collins’ monster”. Despite the lack of name and good documentation, the Collins’ monster have repeatedly featured in studies tackling lobopodian evolution and phylogeny (e.g., Ramsköld & Chen 1998; Budd 2001; Ou et al. 2011; Caron & Aria 2017). It was only in 2020 that the animal was formally named Collinsovermis monstruosus, in honour of Desmond Collins, and fully described based on high resolution pictures of all available material (Caron & Aria 2020).

Description:

Morphology:

Collinsovermis has a plump appearance, with an annulated, unjointed body divided into anterior and posterior regions. The entire dorsum of the body is covered in well-developed spines—three short pairs cover the first three somites (“body segments”) behind the head, while triads of longs spines cover the remaining 10 somites. The anterior region bears 6 pairs of elongate lobopods with thin spines arranged in chevrons, and a small head, as a protrusion bearing a pair of sensory filaments as well as a small dorsal plate and a frontal mouth. The posterior region is made ventrally of 8 pairs of stout annulated lobopods ending in strong, single claws.

Abundance:

Like other lobopodians, Collinsovermis is excessively rare. There are only 3 specimens known, all from the Collins Quarry on Mount Stephen (Fletcher & Collins 2003). They are housed at the Royal Ontario Museum, Department of Natural History.

Maximum Size:
32 mm.

Ecology:

Life habits: Extremely rare
Feeding strategies: Extremely rare
Ecological Interpretations:

Collinsovermis exemplifies suspension-feeding adaptations in lobopodians associated with strongly-developed defensive elements. Like other members of the order Luolishaniidae, and those of the family bearing its name, this animal was using its stout back limbs for anchoring (probably to sponges) and its slender spinose anterior limbs to sieve organic particles or plankton. The long dorsal spines most certainly served as deterrent to predators.

References:

  • BUDD, G. E. 2001. Tardigrades as ‘stem-group arthropods’: The evidence from the Cambrian fauna. Zoologischer Anzeiger, 240, 265–279.
  • CARON, J. and ARIA, C. 2020. The Collins’ monster, a spinous suspension‐feeding lobopodian from the Cambrian Burgess Shale of British Columbia. Palaeontology, 63, 979–994.
  • CARON, J.-B. and ARIA, C. 2017. Cambrian suspension-feeding lobopodians and the early radiation of panarthropods. BMC Evolutionary Biology, 17, 29.
  • DELLE CAVE, L. and SIMONETTA, A. M. 1991. Early Palaeozoic arthropods and problems of arthropod phylogeny; with some notes on taxa of doubtful affinities. In S, S. A. M. C. M. (ed.) The Early Evolution of Metazoa and the Significance of Problematic Taxa. Proceedings of an International Symposium Held at the University of Camerino 27-31 March 1989, Cambridge University Press, 189–244 pp.
  • FLETCHER, T. P. and COLLINS, D. 2003. The Burgess Shale and associated Cambrian formations west of the Fossil Gully Fault Zone on Mount Stephen, British Columbia. Canadian Journal of Earth Sciences, 40, 1823–1838.
  • OU, Q., LIU, J., SHU, D., HAN, J., ZHANG, Z., WAN, X. and LEI, Q. 2011. A rare onychophoran-like lobopodian from the lower Cambrian Chengjiang Lagerstätte. Journal of Paleontology, 85, 587–594.
  • RAMSKÖLD, L. and CHEN, J. Y. 1998. Cambrian lobopodians: morphology and phylogeny. In EDGECOMBE, G. D. (ed.) Arthropod Fossils and Phylogeny, Columbia University Press, New York, 107–150 pp.
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Ovatiovermis cribratus

Artistic reconstruction of Ovatiovermis cribratus. Danielle Dufault © ROM

Taxonomy:

Kingdom: Extremely rare
Phylum: Extremely rare
Higher Taxonomic Assignment: Order Luolishaniida
Species name: Ovatiovermis cribratus
Remarks:

Ovatiovermis is one of a variety of lobopodian taxa from the Cambrian, which are early members of the lineage that gave rise to arthropods, and whose only modern survivors are onychophorans (velvet worms) and tardigrades (water bears). Lobopodians characteristically have annulated, unjointed bodies and bear soft limbs after which they are called: the lobopods. Ovatiovermis groups with Facivermis as a peculiar “unarmoured” lineage within the Order Luolishaniida, notably characterized by their thin spines arranged in chevrons and the differentiation of their body into functional regions for suspension-feeding (Caron & Aria 2017, 2020).

Described by: Caron and Aria
Description date: 2017
Etymology:

Ovatiovermis – from the Latin ovatio (ovation) and vermis (worm) owing to the inferred upward-reaching, limb-waving posture of these animals.
cribratus – from the Latin cribrare, to sieve.

Type Specimens: Holotype ROMIP 52707 and Paratype ROMIP 64006 at the Royal Ontario Museum, Toronto, Canada.
Other species:

Burgess Shale and vicinity: None

Other deposits: None

Age & Localities:

Age:
Middle Cambrian, Wuliuan stage, Bathyuriscus-Elrathina Zone of the Burgess Shale Formation (approximately 507 million years old).
Principal localities:

The Walcott Quarry on Fossil Ridge.

History of Research:

Brief history of research:

Although the first and most complete specimen was discovered in 1994 by one of the ROM expeditions, Ovatiovermis was not described until 2017 by Caron and Aria. The animal served to exemplify suspension-feeding strategies early in panarthropod evolution, and prompted an analysis of the extent and significance of such lifestyle among other lobopodians.

Description:

Morphology:

Ovatiovermis has a worm-like, tubular body divided into three sections: An anterior section with two pairs of very long lobopods bearing thin spines arranges in chevrons and ending in paired claws; a middle section with four pairs of similar but smaller lobopods; and a posterior section with three pairs of very stout lobopods ending in single strong claws. The head is small and bears a pair of simple visual organs and an eversible pharynx. The surface of the body is devoid of any sclerite or spine. Cuticular folds are visible in the space in between lobopods.

Abundance:

Only two specimens. The holotype specimen (ROMIP 52707) was collected in 1994 from bed assemblage −120 (about 1.2 m below the base of the original Walcott Quarry floor), which is distinct from other bed assemblages in that it has particularly well-preserved specimens across a range of taxa (92 species). The paratype specimen (ROMIP 64006) was found in 2016 by a guest on a guided hike to the Walcott Quarry and forwarded to Dr. Jean-Bernard Caron by the Parks Interpreter office.

Maximum Size:
Maximum Size: 25 mm.

Ecology:

Life habits: Extremely rare
Feeding strategies: Extremely rare
Ecological Interpretations:

With its anterior sieving and posterior anchoring appendages, Ovatiovermis exemplifies suspension-feeding specialization among lobopodians. It was likely anchored to sponges and stood erect in the water to catch food particles or plankton.

References:

  • CARON, J. and ARIA, C. 2020. The Collins’ monster, a spinous suspension‐feeding lobopodian from the Cambrian Burgess Shale of British Columbia. Palaeontology, 63, 979–994.
  • CARON, J.-B. and ARIA, C. 2017. Cambrian suspension-feeding lobopodians and the early radiation of panarthropods. BMC Evolutionary Biology, 17, 29.
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Worthenella cambria

Worthenella cambria (USNM 57643) – Holotype, part and counterpart. Left, plate 22 of Walcott (1911), showing a retouched image of the original specimen described (figure 2) together with other “worms.” Right, images of the same specimen. Specimen length = 60 mm. Specimen wet – direct light (left column), dry – polarized light (right column). Walcott Quarry.

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

Taxonomy:

Kingdom: Extremely rare
Phylum: Extremely rare
Higher Taxonomic assignment: Unranked clade (stem group arthropods)
Species name: Worthenella cambria
Remarks:

This animal is related to arthropods, but its systematic status within this group is unknown (Briggs and Conway Morris, 1986).

Described by: Walcott
Description date: 1911
Etymology:

Worthenella – Possibly after the American palaeontologist Amos Henry Worthen, who died in 1888, just as Walcott’s career was taking off.

cambria – from the Welsh Cambria meaning Wales, in reference to the age of the fossil.

Type Specimens: Holotype –USNM57643 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, Bathyuriscus-Elrathina Zone (approximately 505 million years ago).
Principal localities:

The Walcott Quarry on Fossil Ridge.

History of Research:

Brief history of research:

Worthenella was first described by Walcott from a single specimen in a 1911 monograph dealing with various Burgess Shale worms. Walcott interpreted this animal as a polychaete annelid (or bristle worm), in the same family as the animal Wiwaxia (which is now interpreted as a primitive mollusc). However, this interpretation was questioned (Conway Morris, 1979), and the affinities of Worthenella have remained difficult to establish because this singular fossil is too poorly known (Briggs and Conway Morris, 1986).

Description:

Morphology:

The animal is elongate with a small head and bears at least 46 segments of similar dimensions. Appendages or tentacles are present beneath the head, but their preservation is poor and it is difficult to know their precise nature and arrangement. The anterior 34 segments seem to bear filamentous branches on their ventral sides, with the following 8 segments equipped with longer appendages. The gut is straight and the anus is terminal.

Abundance:

This animal is known from a single specimen.

Maximum Size:
60 mm

Ecology:

Life habits: Extremely rare
Feeding strategies: Extremely rare
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.

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

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

Other Links:

None

Waputikia ramosa

3D animation of Waputikia ramosa.

ANIMATION BY PHLESCH BUBBLE © ROYAL ONTARIO MUSEUM

Taxonomy:

Kingdom: Extremely rare
Phylum: Extremely rare
Higher Taxonomic assignment: Non applicable
Species name: Waputikia ramosa
Remarks:

No revisions of this alga have been published since its original description by Walcott (1919) and its affinities remain uncertain.

Described by: Walcott
Description date: 1919
Etymology:

Waputikia – from the Waputik Icefield, a glacier in Yoho National Park, east of the Burgess Shale.

ramosa – from the Latin ramosus, “full of branches,” in reference to the presence of clumps of branches.

Type Specimens: Syntypes –USNM35409, 35410, 35411 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, Bathyuriscus-Elrathina Zone (approximately 505 million years ago).
Principal localities:

The Walcott Quarry on Fossil Ridge.

History of Research:

Brief history of research:

This genus was described by Charles Walcott (1919) as a possible red alga. However, like all the algae from the Burgess Shale, it awaits a modern redescription.

Description:

Morphology:

Waputikia has a large central stem with wide branches at irregular intervals. The large branches divide dichotomously (into two), and the smaller tertiary or quaternary branches divide into much finer branches forming small terminal bush-like structures.

Abundance:

Waputikia is very rare and represents only 0.02% of the Walcott Quarry community (Caron and Jackson, 2008).

Maximum Size:
60 mm

Ecology:

Life habits: Extremely rare
Feeding strategies: Extremely rare
Ecological Interpretations:

No attachment structure for this alga has been preserved but it probably lived attached to the sea floor.

References:

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

WALCOTT, C. 1919. Cambrian Geology and Paleontology IV. Middle Cambrian Algae. Smithsonian Miscellaneous Collections, 67(5): 217-260.

Other Links:

None

Walcottidiscus typicalis

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

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

Taxonomy:

Kingdom: Extremely rare
Phylum: Extremely rare
Higher Taxonomic assignment: Edrioasteroidea (Order: Edrioasteroida, stem group echinoderms)
Species name: Walcottidiscus typicalis
Remarks:

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

Described by: Bassler
Description date: 1935
Etymology:

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

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

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

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

Other deposits: none.

Age & Localities:

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

The Walcott Quarry on Fossil Ridge.

History of Research:

Brief history of research:

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

Description:

Morphology:

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

Abundance:

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

Maximum Size:
64 mm

Ecology:

Life habits: Extremely rare
Feeding strategies: Extremely rare
Ecological Interpretations:

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

References:

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

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

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

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

Other Links:

None

Ulospongiella ancyla

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

© ROYAL ONTARIO MUSEUM. PHOTOS: JEAN-BERNARD CARON

Taxonomy:

Kingdom: Extremely rare
Phylum: Extremely rare
Higher Taxonomic assignment: Demospongea (Order: Monaxonida)
Species name: Ulospongiella ancyla
Remarks:

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

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

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

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

Type Specimens: Holotype –ROM43830 (wrongly referred asROM48830 in Rigby and Collins 2004) in the Royal Ontario Museum, Toronto, Canada.
Other species:

Burgess Shale and vicinity: none.

Other deposits: none.

Age & Localities:

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:

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

Description:

Morphology:

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

Abundance:

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

Maximum Size:
19 mm

Ecology:

Life habits: Extremely rare
Feeding strategies: Extremely rare
Ecological Interpretations:

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

References:

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

Other Links:

None

Thelxiope palaeothalassia

Thelxiope palaeothalassia (GSC 74990). Articulated specimen (close up to the right), associated with several individuals of the arthropod Canadaspis perfecta. Specimen length = 29 mm. Specimen dry – polarized light. Walcott Quarry.

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

Taxonomy:

Kingdom: Extremely rare
Phylum: Extremely rare
Higher Taxonomic assignment: Unranked clade (stem group arthropods)
Species name: Thelxiope palaeothalassia
Remarks:

The affinity of Thelxiope has not been considered in detail because the appendages are unknown.

Described by: Simonetta and Delle Cave
Description date: 1975
Etymology:

Thelxiope – from the Greek thelx meaning “enchanting,” and ops, meaning “voice,” referring to the muse-like appearance of the animal.

palaeothalassia – from the Greek palaios, meaning “ancient,” and thalassios, meaning “marine,” in reference to the age and environment where the animal lived.

Type Specimens: Holotype –USNM144914 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, Bathyuriscus-Elrathina Zone (approximately 505 million years ago).
Principal localities:

The Walcott Quarry on Fossil Ridge.

History of Research:

Brief history of research:

Walcott (1912) figured two fragmentary specimens as Mollisoniarara; these were first reinterpreted by Simonetta (1964) within a new genus Parahabelia rara, along with three additional specimens that he thought were related. However, Simonetta and Delle Cave (1975) considered that among those five specimens, the two originally figured by Walcott as M? rara had to be synonymized with M. symmetrica and the other three had to be placed within a new genus and species called Thelxiope palaeothalassia, a name in use since then.

Description:

Morphology:

This species has a relatively wide cephalon and seven segments and resembles Habelia in overall shape. However, in T. palaeothalassia, each segment bears a single prominent spine pointing dorsally. The last segment is armed with a very long pointed telson.

Abundance:

Thelxiope is extremely rare, with only four known specimens.

Maximum Size:
43 mm

Ecology:

Life habits: Extremely rare
Feeding strategies: Extremely rare
Ecological Interpretations:

Thelxiope is too poorly known to allow detailed studies of its ecology.

References:

SIMONETTA, A. M. 1964. Osservazioni sugli arthropodi non trilobiti della “Burgess Shale” (Cambriano medio). Monitore Zoologico Italiano, 72 (3-4: III Contributo: I Generi MolariaHabeliaEmeraldellaParahabelia (Nov.) Emeraldoides (Nov.): 215-231.

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

WALCOTT, C. 1912. Cambrian Geology and Paleontology II. Middle Cambrian Branchiopoda, Malacostraca, Trilobita and Merostomata. Smithsonian Miscellaneous Collections, 57(6):145-228.

Other Links:

None

Thaumaptilon walcotti

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

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

Taxonomy:

Kingdom: Extremely rare
Phylum: Extremely rare
Higher Taxonomic assignment: Unranked clade (stem group cnidarians)
Species name: Thaumaptilon walcotti
Remarks:

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

Described by: Conway Morris
Description date: 1993
Etymology:

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

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

Type Specimens: Holotype –USNM468028 in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.
Other species:

Burgess Shale and vicinity: none.

Other deposits: none.

Age & Localities:

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

The Walcott Quarry on Fossil Ridge.

History of Research:

Brief history of research:

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

Description:

Morphology:

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

Abundance:

Only three specimens are known.

Maximum Size:
210 mm

Ecology:

Life habits: Extremely rare
Feeding strategies: Extremely rare
Ecological Interpretations:

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

References:

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

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

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

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

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

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

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

Other Links:

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

Tegopelte gigas

Tegopelte gigas (USNM 189201) – Holotype. Complete specimen showing antennae and appendages partially prepared near the back. Specimen length = 270 mm. Specimen dry – direct (top) and polarized light (bottom). Walcott Quarry.

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

Taxonomy:

Kingdom: Extremely rare
Phylum: Extremely rare
Higher Taxonomic assignment: Unranked clade (stem group arthropods)
Species name: Tegopelte gigas
Remarks:

Tegopelte is usually compared to the soft-bodied “trilobites” such as Naraoia and Saperion, but the exact relationships of these taxa to the mineralized trilobites is uncertain (Whittington, 1977). The tegopeltids and other trilobite-like arthropods are sometimes referred to as Trilobitoidea, which when grouped together with the trilobites form the Lamellipedians (Hou and Bergström, 1997; Wills et al., 1998; Edgecombe and Ramsköld, 1999). This group has been variously placed in the upper stem lineage of the arthropods (Budd, 2002), or in the stem lineage of either the mandibulates (Scholtz and Edgecombe, 2006) or the chelicerates (Cotton and Braddy, 2004).

Described by: Simonetta and Delle Cave
Description date: 1975
Etymology:

Tegopelte – from the Greek tegos, “tile,” and pelte, “leather-shield,” referring to the shape of the dorsal body covering.

gigas – from the Greek gigas, “giant,” referring to the large size of the animal.

Type Specimens: Holotype –USNM189201 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, Bathyuriscus-Elrathina Zone (approximately 505 million years ago).
Principal localities:

The Walcott Quarry on Fossil Ridge.

History of Research:

Brief history of research:

Tegopelte was first described by Simonetta and Delle Cave (1975) to include only two relatively large specimens. This original description showed Tegopelte to have a cephalon with six or seven pairs of walking appendages, a thorax of four tergites each bearing five appendages, and a tail segment with ten appendages. Whittington (1985) re-examined the animal, reducing the number of head appendages to three, and describing the thorax as having only three tergites with three appendages each. The tail in Whittington’s (1985) reconstruction had two segments with a total of 20 appendages. Later re-examination by Ramsköld et al. (1996) suggested that the body has no tergites, but instead consists of an undivided dorsal shield. Tegopelte has been grouped together with the Chengjiang taxon Saperion to form the Tegopeltidae (Ramsköld et al., 1996; Hou and Bergström, 1997), a clade later confirmed by cladistic analysis (Edgecombe and Ramsköld, 1999; Hendricks and Lieberman, 2008).

Description:

Morphology:

The dorsal morphology of Tegopelte consists of an elongated oval-shaped dorsal shield that is featureless and undivided. The length of the two known specimens is 25.7 cm and 27.0 cm, making it one of the largest arthropods in the Burgess Shale. The ventral morphology consists of a pair of multi-segmented antennae at the front of the body, followed by a series of identical limbs that are segmented and branch into two (biramous), totaling approximately 33 along the entire body. The biramous limbs have a walking branch made up of six segments with a pair of spines on the terminal segment, and a filamentous branch where numerous elongated oval blades attach to a central shaft. The biramous limbs decrease in size towards the posterior end of the body.

Abundance:

Tegopelte is extremely rare, with only two known specimens.

Maximum Size:
270 mm

Ecology:

Life habits: Extremely rare
Feeding strategies: Extremely rare
Ecological Interpretations:

Tegopelte probably spent much of its time walking on the seafloor, based on the presence of many appendages. It used the segmented branches of its biramous appendages for walking, and it is likely that the filamentous branches were used for oxygen exchange, and to propel the animal through the water during short bursts of swimming. The antennae would have been used to sense the environment. The lack of eyes, gut glands and feeding appendages make it difficult to allocate a feeding strategy to Tegopelte.

References:

BUDD, G. E. 2002. A palaeontological solution to the arthropod head problem. Nature, 417: 271-275.

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

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.

EDGECOMBE, G. D. and L. RAMSKÖLD. 1999. Relationships of Cambrian Arachnata and the systematic position of Trilobita. Jounral of Paleontology, 73: 263-287.

HENDRICKS, J. R. AND B. S. LIEBERMAN. 2008. New phylogenetic insights into the Cambrian radiation of arachnomorph arthropods. Journal of Paleontology, 83: 585-594.

HOU, X. AND J. BERGSTRÖM. 1997. Arthropods of the Lower Cambrian Chengjiang fauna, southwest China. Fossils and Strata, 45: 1-116.

RAMSKÖLD, L., J. CHEN, G. D. EDGECOMBE AND G. ZHOU. 1996. Preservational folds simulating tergite junctions in tegopeltid and naraoiid arthropods. Lethaia, 29: 15-20.

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

WHITTINGTON, H. B. 1977. The Middle Cambrian trilobite Naraoia, Burgess Shale, British Columbia. Philosophical Transactions of the Royal Society of London, B, 280: 409-443.

WHITTINGTON, H. B. 1985. Tegopelte gigas, a second soft-bodied trilobite from the Burgess Shale, Middle Cambrian, British Columbia. Journal of Paleontology, 59: 1251-1274.

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

Other Links:

None