The Burgess Shale

Phylas: Onychophora (Lobopodia)

Collinsovermis monstruosus

Collinsovermis monstruosus, holotype ROMIP 52703

Taxonomy:

Kingdom: Onychophora (Lobopodia)
Phylum: Onychophora (Lobopodia)
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: Onychophora (Lobopodia)
Feeding strategies: Onychophora (Lobopodia)
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.
Other Links:

Ovatiovermis cribratus

Artistic reconstruction of Ovatiovermis cribratus. Danielle Dufault © ROM

Taxonomy:

Kingdom: Onychophora (Lobopodia)
Phylum: Onychophora (Lobopodia)
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: Onychophora (Lobopodia)
Feeding strategies: Onychophora (Lobopodia)
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.
Other Links:

Hallucigenia sparsa

3D animation of Hallucigenia sparsa.

Animation by Phlesch Bubble © Royal Ontario Museum

Taxonomy:

Kingdom: Onychophora (Lobopodia)
Phylum: Onychophora (Lobopodia)
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: Onychophora (Lobopodia)
Feeding strategies: Onychophora (Lobopodia)
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

Aysheaia pedunculata

Reconstruction of Aysheaia pedunculata.

© Marianne Collins

Taxonomy:

Kingdom: Onychophora (Lobopodia)
Phylum: Onychophora (Lobopodia)
Higher Taxonomic assignment: Xenusia (Order: Scleronychophora, stem group onychophorans)
Species name: Aysheaia pedunculata
Remarks:

Aysheaia 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:

Aysheaia – after the nearby Aysha peak (since renamed Ayesha peak) in the Wapta icefield (3,065 m); original meaning unknown.

pedunculata – from the Latin pedunculus, “foot.”

Type Specimens: Holotype –USNM57655 in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.
Other species:

Burgess Shale and vicinity: none

Other deposits: A.? prolata from the Middle Cambrian of Utah (Robison, 1985).

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 originally described Aysheaia as an annelid worm (Walcott, 1911). It was later re-described as a velvet worm (or a close relative) (Brues, 1923; Hutchinson, 1930; Walcott, 1931; Walton, 1927), although it lacked features such as jaws and slime glands. Its position remains a subject of debate, with a position in a new phylum being mooted (Tiegs and Manton, 1958). A morphological reinterpretation based on photographs (Delle Cave and Simonetta, 1975) prompted a detailed re-study of the fossil specimens (Whittington, 1978), and relationships were suggested with the water bears (tardigrades) (Bergström, 1978). Aysheaia is now grouped with close relatives in the class Xenusia (Liu et al., 2008), lobopods that fall on the arthropod stem lineage (Budd, 1996, 1998; Whittington, 1978).

Description:

Morphology:

Aysheaia is a worm-like animal, 1 to 6 cm in length and about 5 mm broad, bearing ten pairs of clawed, spiny limbs on the lower part of its body. It did not have a separate head, but a mouth occupied the very front of the body, accompanied by a pair of appendages and a circlet of bumps (papillae). The animal had a soft, flexible, non-mineralized cuticle, which had a corrugated, accordion-like form. Each stubby limb had ten corrugations, some of which bore a spiny projection. A suite of claws also adorned the end of each stub-foot. A faint line running down the axis of the organism is interpreted as its gut.

Abundance:

Aysheaia is rare in the Walcott Quarry representing less than 0.04% of the specimens counted in the community (Caron and Jackson, 2008).

Maximum Size:
60 mm

Ecology:

Life habits: Onychophora (Lobopodia)
Feeding strategies: Onychophora (Lobopodia)
Ecological Interpretations:

Aysheaia is frequently associated with the remains of sponges, and an ecological association has been posited. Whether Aysheaia used its spines to adhere to sponges while feeding on them, or whether it simply hid among sponges for protection from predators, is unclear.

References:

BERGSTRÖM, J. 1978. Morphology of fossil arthropods as a guide to phylogenetic relationships, p. 1-56 In A. P. Gupta (ed.), Arthropod Phylogeny. Van Nostrand Reinhold Co. New York.

BRUES, C. T. 1923. The geographical distribution of the Onychophora. American Naturalist, 57: 210-217.

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

BUDD, G. E. 1998. Stem group arthropods from the Lower Cambrian Sirius Passet fauna of North Greenland, p. 125-138. In R. A. Fortey and R. H. Thomas (eds.), Arthropod relationships. Volume 55. Chapman & Hall, London.

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

DELLE CAVE, L. AND A. M. SIMONETTA. 1975. Notes on the morphology and taxonomic position of Aysheaia (Onycophora?) and of Skania (undetermined phylum). Monitore Zoologico Italiano, 9: 67-81.

HUTCHINSON, G. E. 1930. Restudy of some Burgess Shale fossils. Proceedings of the United States National Museum, 78(11): 59.

LIU, J., D. SHU, J. HAN, Z. ZHANG, AND X. ZHANG. 2008. Origin, diversification, and relationships of Cambrian lobopods. Gondwana Research, 14(1-2): 277-283.

ROBISON, R. A. 1985. Affinities of Aysheaia (Onychophora), with description of a new Cambrian species. Journal of Paleontology, 59(1): 226-235.

TIEGS, O. W. AND S. M. MANTON. 1958. The evolution of the Arthropoda. Biological Reviews, 33(3): 255-333.

WALCOTT, C. D. 1911. Middle Cambrian annelids. Smithsonian Miscellaneous Collections, 57(2): 109-144.

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

WALTON, L. B. 1927. The polychaete ancestry of the insects. American Naturalist, 61: 226-250.

WHITTINGTON, H. B. 1978. The lobopod animal Aysheaia pedunculata Walcott, Middle Cambrian, Burgess Shale, British Columbia. Philosophical Transactions of the Royal Society of London. B, Biological Sciences, 284(1000): 165-197.

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:

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