Media: 2D Model



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Pakucaris apatis

Artistic reconstruction of Pakucaris apatis. Danielle Dufault © ROM

Taxonomy:

Kingdom: 2D Model
Phylum: 2D Model
Higher Taxonomic assignment: Hymenocarines, Family: Odaraiidae
Species name: Pakucaris apatis
Remarks:

Hymenocarines were early arthropods with bivalved carapaces and mandibles, forming the bulk of the first mandibulates (represented today by myriapods, crustaceans and insects) (Aria and Caron 2017; Vannier et al. 2018). In many hymenocarines, including Pakucaris, determining the exact number and types of appendages in their head remains difficult, which hinders a detailed understanding of the evolutionary relationships inside this group. Pakucaris most probably belongs to the family Odaraiidae, a group of hymenocarines with highly multisegmented bodies, reduced or absent antennae and highly multisegmented legs.

Described by: Izquierdo-López & Caron
Description date: 2021
Etymology:

Pakucaris – from the Japanese onomatopoeia paku, suggestive of ‘eating’, related to the video game character Pac-Man, due to the naked eye resemblance of the carapace and shield of Pakucaris to the shape of the character. Latin caris, meaning “crab” or “shrimp”, and

apatis – from the goddess of deception in Greek mythology Apate, in reference to the resemblance of Pakucaris to a trilobite.

Type Specimens: Holotype ROMIP65739
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, Tokumm Creek

History of Research:

Brief history of research:

The holotype of Pakucaris apatis was first discovered during the 2012 expedition to the Marble Canyon site of the Burgess Shale. A few other specimens were discovered during the following 2014 and 2016 expeditions and classified as “New arthropod E” (Nanglu et al. 2020). The 2018 expedition at the Tokumm Creek site uncovered one additional specimen. The first description of Pakucaris apatis was published in 2021 in the journal Papers in Paleontology (Izquierdo-López and Caron, 2021). Several other authors have noted the similarity between the shield of Pakucaris and pygidia (O’Flynn et al. 2022). A pygidium is a structure in which the most posterior segments of an arthropod become fused, usually into a shield. The pygidium is typically found in trilobites, but also across many other groups in the Cambrian, suggesting that this structure appeared multiple times independently.

Description:

Morphology:

Pakucaris has two morphotypes: a small one (around 1 cm) with its body subdivided into 30-35 segments, and a larger one (around 2.5 cm), with its body subdivided into 70-80 segments. The carapace of Pakucaris covers up to two-thirds of the total body length. It has a dome-like shape with a small dorsal crest that runs across its entire length. The carapace bends towards the front, extending into a small process (rostrum). Similarly, the lateral sides of the carapace also extend frontally into small lateral processes. The head has one pair of pedunculate eyes, one pair of thin small appendages, and at least one pair of larger segmented antennae. The small thin appendages are not segmented and represent a sensorial organ known as frontal filaments. The first antennae (also termed antennules) have 7 to 8 segments, with each segment bearing a small spine. Each segment of the body bears one pair of limbs, each subdivided into two branches (biramous): a walking leg (endopod) and a paddle-like flap (exopod). The endopod is thin and is subdivided into at least 20-21 segments. The exopod has an ovoid, flattened shape and is as long as half the endopod. The posterior section of the body has a shield-like structure. This shield is formed by the fusion and lateral extensions of the segments. The shield bears around 10 big spines on each of its lateral sides, as well as a series of smaller spines on its posterior side.

Abundance:

Pakucaris is rare, only known from eight specimens from the Marble Canyon and Tokumm Creek sites. The bigger morphotype is only known from one specimen.

Maximum Size:
About 2.5 cm

Ecology:

Life habits: 2D Model
Feeding strategies: 2D Model
Ecological Interpretations:

Pakucaris was probably a nektobenthic animal living close to the benthos (Izquierdo-López and Caron 2021). It may have used its antennae with spines to scrape rocks or other objects and may have also used its paddle-like exopods to create currents and capture organic particles, aided by its antennae and other head appendages. The tail shield (or pygidium) of Pakucaris was most probably a structure to protect against predators. The two morphotypes of Pakucaris may represent different growth stages of males and females, but the number of specimens available to date is too limited to reach a conclusion.

References:

  • ARIA, C. and CARON, J. B. 2017. Burgess Shale fossils illustrate the origin of the mandibulate body plan. Nature, 545: 89–92.
  • IZQUIERDO-LÓPEZ, A. and CARON, J. B. 2021. A Burgess Shale mandibulate arthropod with a pygidium: a case of convergent evolution. Papers in Palaeontology, 7: 1877–1894.
  • NANGLU, K., CARON, J. and GAINES, R. 2020. The Burgess Shale paleocommunity with new insights from Marble Canyon, British Columbia. Paleobiology, 46(1): 58–81.
  • O’FLYNN, R. J., WILLIAMS, M., YU, M., HARVEY, T. and LIU, Y. 2022. A new euarthropod with large frontal appendages from the early Cambrian Chengjiang biota. Palaeontologia Electronica, 25(1):a6: 1–21.
  • VANNIER, J., ARIA, C., TAYLOR, R. S. and CARON, J. B. 2018. Waptia fieldensis Walcott, a mandibulate arthropod from the middle Cambrian Burgess Shale. Royal Society Open Science, 5:172206.
Other Links:


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Acanthotretella spinosa

Reconstruction of Acanthotretella spinosa.

© MARIANNE COLLINS

Taxonomy:

Kingdom: 2D Model
Phylum: 2D Model
Higher Taxonomic assignment: Lingulata (Order: Siphonotretida, stem group brachiopods)
Species name: Acanthotretella spinosa
Remarks:

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

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

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

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

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

Burgess Shale and vicinity: none.

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

Age & Localities:

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:

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

Description:

Morphology:

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

Abundance:

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

Maximum Size:
8 mm

Ecology:

Life habits: 2D Model
Feeding strategies: 2D Model
Ecological Interpretations:

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

References:

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

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

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

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

Other Links:

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



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Yohoia tenuis

3D animation of Yohoia tenuis.

ANIMATION BY PHLESCH BUBBLE © ROYAL ONTARIO MUSEUM

Taxonomy:

Kingdom: 2D Model
Phylum: 2D Model
Higher Taxonomic assignment: Megacheirans, Clade Cheiromorpha
Species name: Yohoia tenuis
Remarks:

Henriksen (1928) created the family Yohoiidae, and later Simonetta and Delle Cave (1975) erected the order Yohoiida—however, they have so far remained monogeneric (no other included genus other than Yohoia), and are therefore not taxonomically meaningful. Yohoia was originally considered to be a branchiopod crustacean (Walcott, 1912; Simonetta, 1970), but it is now understood as an iconic representative of the class Megacheira. Megacheirans are basal true arthropods with a frontal appendage pointing upward and made of multiple claws (the cheira, or “great appendage”). The relationship of megacheirans with respect to extant lineages is still debated, but cumulative evidence has recently favoured a closer affinity with chelicerates (Aria, 2022).

Described by: Walcott
Description date: 1912
Etymology:

Yohoia – from the Yoho River, Lake, Pass, Glacier, Peak (2,760 m) and Park, British Columbia, Canada. “Yoho” is a Cree word expressing astonishment.

tenuis – from the Latin tenuis, “thin,” referring to its slender body.

Type Specimens: Lectoype –USNM57699 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, Wuliuan stage, Burgess Shale Formation (approximately 505 million years ago).
Principal localities:

The Walcott, Raymond and Collins Quarries on Fossil Ridge.

History of Research:

Brief history of research:

Yohoia was first described by Walcott (1912), who designated the type species Y. tenuis based on six specimens, and a second species, Y. plena, based on one specimen. Additional specimens of Y. tenuis were described by Simonetta (1970), and a major redescription of Yohoia tenuis was then undertaken by Whittington (1974), based on over 400 specimens of this species. Whittington (1974) invalidated Y. plena, upgrading it to its own genus, Plenocaris plena, leaving Y. tenuis as the only species of Yohoia. Another thorough revision was published much later by Haug et al. (2012), who specifically compared the raptorial appendages of Yohoia with chelicerae. Yohoia appears to represent an evolutionary intermediate between early megacheiran taxa called jianfengiids and the relatives of Leanchoilia (Aria et al., 2020).

Description:

Morphology:

The body of Yohoia consists of a head region encapsulated in a cephalic shield and 13 trunk segments, ending in a paddle-shaped telson. The dorsal head shield is roughly square and extends over the dorsal and lateral regions of the head. There is a pair of great appendages, or cheirae, at the front of the head. Each appendage consists of two long segments that bend like an elbow at their articulation, with four long spines at the tip. Three pairs of biramous appendages with walking and increasingly long swimming branches project from beneath the head shield behind the great appendages. The body behind the head consists of ten segments that extend over the back and down the side of the animal, ending in backward-facing triangular points. These trunk segments bear appendages similar to those in the head. The last three body segments have no appendages, and the telson is a paddle-shaped plate with distal spines.

Abundance:

Over 700 specimens of Yohoia are known from the Walcott Quarry, comprising 1.3% of the specimens counted (Caron and Jackson, 2008) but only few specimens are known from the Raymond and Collins Quarries.

Maximum Size:
23 mm

Ecology:

Life habits: 2D Model
Feeding strategies: 2D Model
Ecological Interpretations:

Like other megacheirans, and a number of modern crustaceans, Yohoia was probably living close to the sea floor and using its raptorial appendages to prey on small animals. Prey items were most likely non-biomineralized, owing to the lack of strong masticatory devices. Breathing could have taken place through the thin, paddle-shaped branches of the appendages, but it is also possible that Yohoia shared dedicated gills with other megacheirans (Liu et al., 2021; Aria et al., 2023).

References:

  • ARIA, C. (2022) The origin and early evolution of arthropods. Biological Reviews 97, 1786–1809.
  • ARIA, C., VANNIER, J., PARK, T.S. & GAINES, R.R. (2023) Interpreting fossilized nervous tissues. BioEssays, 2200167.
  • ARIA, C., ZHAO, F., ZENG, H., GUO, J. & ZHU, M. (2020) Fossils from South China redefine the ancestral euarthropod body plan. BMC Evolutionary Biology 20, 4.
  • HAUG, J.T., WALOSZEK, D., MAAS, A., LIU, Y. & HAUG, C. (2012) Functional morphology, ontogeny and evolution of mantis shrimp-like predators in the Cambrian. Palaeontology 55, 369–399.
  • HENRIKSEN, K.L. (1928) Critical notes upon some Cambrian arthropods described by Charles D. Walcott. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening: Khobenhavn 86, 1–20.
  • LIU, Y., EDGECOMBE, G.D., SCHMIDT, M., BOND, A.D., MELZER, R.R., ZHAI, D., MAI, H., ZHANG, M. & HOU, X. (2021) Exites in Cambrian arthropods and homology of arthropod limb branches. Nature Communications 12, 4619.
  • SIMONETTA, A.M. (1970) Studies on non trilobite arthropods of the Burgess Shale (Middle Cambrian). Palaeontographia Italica 66 (New series 36), 35–45.
  • SIMONETTA, A.M. & DELLE CAVE, L. (1975) The Cambrian non trilobite arthropods from the Burgess Shale of British Columbia. A study of their comparative morphology taxinomy 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.
  • WHITTINGTON, H.B. (1974) Yohoia Walcott and Plenocaris n. gen., arthropods from the Burgess Shale, Middle Cambrian, British Columbia. Geological Survey of Canada Bulletin, Department of Energy, Mines and Resources Canada 231, 1–21.
Other Links:

None



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Wiwaxia corrugata

3D animation of Wiwaxia corrugata grazing on Morania confluens.

ANIMATION BY PHLESCH BUBBLE © ROYAL ONTARIO MUSEUM

Taxonomy:

Kingdom: 2D Model
Phylum: 2D Model
Higher Taxonomic assignment: Unranked clade halwaxiids (stem group molluscs)
Species name: Wiwaxia corrugata
Remarks:

The relationship of Wiwaxia is hotly debated; its similarities to the molluscs have been highlighted (Conway Morris, 1985; Scheltema et al., 2003; Caron et al., 2006; Caron et al., 2007), but Matthew’s original view that it was related to the annelid worms (Matthew, 1899) still finds some adherents (Butterfield, 1990; Conway Morris and Peel, 1995; Butterfield, 2006; 2008). It is also possible that Wiwaxia branched off before the molluscs and annelids diverged (Eibye-Jacobsen, 2004). Wiwaxia has recently been placed in a group called the halwaxiids, along with the halkieriids, Orthrozanclus, and Odontogriphus (Conway Morris and Caron, 2007).

Described by: Matthew
Description date: 1899
Etymology:

Wiwaxia – from Wiwaxy Peaks (2,703 m) in Yoho National Park. The word wiwaxy is originally from the Stoney First Nation Nakoda language, meaning “windy.”

corrugata – from the Latin corrugis, “folded, or wrinkled,” in reference to the wrinkled aspect of the sclerites.

Type Specimens: Holotype –ROM8596 in the Royal Ontario Museum, Toronto, Canada.
Other species:

Burgess Shale and vicinity: none.

Other deposits: none described, although sclerites have been reported from a number of Middle Cambrian deposits extending from northern Canada (Butterfield, 1994) to China (Zhao et al., 1994).

Age & Localities:

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

The Walcott and Raymond Quarries on Fossil Ridge. The Trilobite Beds, Tulip Beds (S7) and Collins Quarry on Mount Stephen. Additional smaller localities are known on Mount Field and Mount Odaray.

History of Research:

Brief history of research:

In an early review of fossils collected from the Trilobite Beds on Mount Stephen by Walker, Canadian palaeontologist G. F. Matthew (1899) described several forms he thought represented tubes of various annelid worms, including one he named Orthotheca corrugata. At the time, Matthew did not know this particular fossil was only part of a much larger organism. It was only when Walcott (1911) discovered articulated and much better preserved specimens from the Phyllopod Bed that the morphology of this species became clearer. Walcott placed corrugata in his new genus Wiwaxia and interpreted it as a polychaete annelid worm (Walcott, 1911). The single best specimen of Walker’s “Orthotheca corrugata” remained unrecognized until it was “rediscovered” in the ROM collections in 1977.

Walcott’s interpretation was called into question in a comprehensive reassessment of the genus (Conway Morris, 1985), and Conway Morris’s link between Wiwaxia mouthparts and the molluscan radula was built upon by Scheltema et al. (2003) and Caron et al. (2006). Butterfield (1990), however, defended an annelid affinity mostly based on the study of individual sclerites, first at the crown-, and later at the stem-group level (Butterfield, 2003; 2006), but further work suggested that the evidence does not conclusively support a close relationship with annelids (Eibye-Jacobsen, 2004). A connection with the halkieriids was drawn early on (Bengtson and Morris, 1984; Conway Morris and Peel, 1995), and expanded more recently (Conway Morris and Caron, 2007).

Other studies have dealt more specifically with the ecology and taphonomy of this animal. The finely spaced patterning of ridges on the scale may have given Wiwaxia an iridescent aspect in life (Parker, 1998). Wiwaxia has proven useful in calculating the extent of decay in fossil assemblages (Caron and Jackson, 2006) and in reconstructing the longer term taphonomic processes responsible for the preservation of the Burgess Shale fossils (Butterfield et al., 2007).

Description:

Morphology:

Wiwaxia corrugata is a slug-like organism up to 5.5 cm in length almost entirely covered (except on the ventral surface) with an array of scale-like elements referred to as sclerites and spines. The body is roughly oval, and lacks evidence of segmentation. The body-covering sclerites are arranged in about 50 rows. In addition, two rows of 7–11 blade-like spines are present on the dorsal surface. Spines and sclerites were inserted directly into the body wall. Wiwaxia’s feeding apparatus consists of two (in rare cases three) toothed plates that have been compared to a molluscan radula or annelid jaws.

Abundance:

Wiwaxia is mostly known from the Walcott Quarry where it is relatively common, representing 0.9% of the specimens counted in the community (Caron and Jackson, 2008).

Maximum Size:
55 mm

Ecology:

Life habits: 2D Model
Feeding strategies: 2D Model
Ecological Interpretations:

The similarity of Wiwaxia’s feeding apparatus to that of Odontogriphus suggests that it too fed on the cyanobacterial Morania mats growing on the Cambrian sea floor. Its sclerite armour-plating and long spines, sometimes found broken, suggest that it was targeted by unidentified predators.

References:

BENGSTON, S. AND S. CONWAY MORRIS, 1984. A comparative study of Lower Cambrian Halkieria and Middle Cambrian Wiwaxia. Lethaia, 17:307-329.

BUTTERFIELD, N. J. 1990. A reassessment of the enigmatic Burgess Shale fossil Wiwaxia corrugata (Matthew) and its relationship to the polychaete Canadia spinosa Walcott. Paleobiology: 287-303.

BUTTERFIELD, N. J. 1994. Burgess Shale-type fossils from a Lower Cambrian shallow-shelf sequence in northwestern Canada. Nature, 369(6480): 477-479.

BUTTERFIELD, N. J. 2003. Exceptional fossil preservation and the Cambrian Explosion. Integrative and Comparative Biology, 43:166-177.

BUTTERFIELD, N. J. 2006. Hooking some stem-group “worms”: fossil lophotrochozoans in the Burgess Shale. BioEssays, 28: 1161-1166.

BUTTERFIELD, N. J. 2008. An early Cambrian radula. Journal of Paleontology, 82(3): 543-554.

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

CARON, J.-B., A. H. SCHELTEMA, C. SCHANDER AND D. RUDKIN, 2006. A soft-bodied mollusc with radula from the Middle Cambrian Burgess Shale. Nature, 442(7099): 159-163.

CARON, J.-B., A. H. SCHELTEMA, C. SCHANDER AND D. RUDKIN. 2007. Reply to Butterfield on stem-group “worms:” fossil lophotrochozoans in the Burgess Shale. BioEssays, 29:200-202.

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(1134): 507-582.

CONWAY MORRIS, S. AND J.-B. CARON, 2007. Halwaxiids and the Early Evolution of the Lophotrochozoans. Science, 315(5816): 1255-1258.

CONWAY MORRIS, S. AND J. S. PEEL, 1995. Articulated halkieriids from the Lower Cambrian of North Greenland and their role in early protostome evolution. Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences, 347(1321): 305-358.

EIBYE-JACOBSEN, D. 2004. A reevaluation of Wiwaxia and the polychaetes of the Burgess Shale. Lethaia, 37(3): 317-335.

MATTHEW, G. F. 1899. Studies on Cambrian Faunas, No. 3. Upper Cambrian fauna, Mount Stephen, British Columbia. The trilobites and worms. Transactions of the Royal Society, 5: 39-66.

PARKER, A. R. 1998. Colour in Burgess Shale animals and the effect of light on evolution in the Cambrian. Proceedings of the Royal Society B: Biological Sciences, 265(1400): 967.

SCHELTEMA, A. H., K. KERTH AND A. M. KUZIRIAN, 2003. Original molluscan radula: Comparisons among Aplacophora, Polyplacophora, Gastropoda, and the Cambrian fossil Wiwaxia corrugata. Journal of Morphology, 257(2): 219-245.

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

ZHAO, Y.-l., Y. QIAN AND X.-S. LI, 1994. Wiwaxia from Early-Middle Cambrian Kaili Formation in Taijiang, Guizhou. Acta Palaeontologica Sinica, 33:359-366.

Other Links:

http://www.paleobiology.si.edu/burgess/wiwaxia.html



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Sidneyia inexpectans

3D animation of Sidneyia inexpectans.

ANIMATION BY PHLESCH BUBBLE © ROYAL ONTARIO MUSEUM

Taxonomy:

Kingdom: 2D Model
Phylum: 2D Model
Higher Taxonomic assignment: Unranked clade (stem group arthropods)
Species name: Sidneyia inexpectans
Remarks:

Sidneyia is usually considered to be closely related to the chelicerates, but its exact position relative to this group remains unclear (Budd and Telford, 2009). Sidneyia has been variously placed as the sister group to the chelicerates (Hou and Bergström, 1997), close to the crown on the chelicerate stem lineage (Bruton, 1981; Edgecombe and Ramsköld, 1999; Hendricks and Lieberman, 2008), or basal in the chelicerate stem lineage (Briggs and Fortey, 1989; Wills et al., 1998; Cotton and Braddy, 2004).

Described by: Walcott
Description date: 1911
Etymology:

Sidneyia – after Walcott’s son Sidney, who discovered the first specimen in August of 1910.

inexpectans – from the Latin inexpectans, “unexpected,” since Walcott did not expect to find such a fossil in strata older than the Ordovician.

Type Specimens: Lectotype –USNM57487 (S. inexpectans) in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.
Other species:

Burgess Shale and vicinity: none.

Other deposits: A single specimen from the Chengjiang Fauna in China was used to describe a second species, Sidneyia sinica (Zhang et al. 2002), however this was later shown to be incorrectly attributed to Sidneyia (Briggs et al. 2008).

Age & Localities:

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

Burgess Shale and vicinity: The Walcott, Raymond and Collins Quarries on Fossil Ridge, Mount Field and Mount Stephen – Tulip Beds (S7) and other smaller localities – Odaray Mountain and Stanley Glacier.

Other deposits: Sidneyia has been described from the Wheeler Formation (Briggs and Robison, 1984) and the Spence Shale (Briggs et al. 2008) in Utah, and the Kinzers Formation in Pennsylvania (Resser and Howell, 1938).

History of Research:

Brief history of research:

Sidneyia was the first fossil to be described by Walcott (1911) from the Burgess Shale. Further details were added by Walcott the following year (Walcott, 1912), and Strømer (1944) and Simonetta (1963) made minor revisions to Walcott’s reconstruction. A large appendage found in isolation was originally suggested to be the large frontal appendage of Sidneyia (Walcott, 1911), but this was later found to belong to the anomalocaridid Laggania (Whittington and Briggs, 1985). A major study by Bruton (1981) redescribed the species based on the hundreds of available specimens.

Description:

Morphology:

Sidneyia has a short, wide head shield that is convexly domed and roughly square. The two front lateral corners are notched to allow an antenna and a stalked eye to protrude. Other than the pair of antennae, which are long and thin with at least 20 segments, there are no cephalic appendages. The hemispherical and highly reflective eyes are above and posterior to the antennae.

The thorax of Sidneyia has nine wide, thin body segments that widen from the first to the fourth segment and then get progressively narrower posteriorly. The first four thoracic segments bear appendages with a large, spiny basal segment (the coxa) and 8 thinner segments, ending in a sharp claw. The next five thoracic appendages have a similar appendage but also have flap-like filaments in association with the limbs.

The abdomen consists of three circular rings that are much narrower than the thorax, with a terminal, triangular telson. The last segment of the abdomen has a pair of wide flaps that articulate with the telson to form a tail fan. A trace of the straight gut can be seen in some specimens extending from the anterior mouth to the anus on the telson, and pieces of broken trilobites are sometimes preserved in the gut.

Abundance:

Sidneyia is a relatively common arthropod in the Walcott Quarry, comprising 0.3% of the specimens counted (Caron and Jackson, 2008). Hundreds of specimens have been collected from the Walcott Quarry (Bruton, 1981) and in other nearby localities.

Maximum Size:
160 mm

Ecology:

Life habits: 2D Model
Feeding strategies: 2D Model
Ecological Interpretations:

Sidneyia walked and swam above the sea floor. Its anterior four thoracic appendages were used for walking, and the spiny basal coxa would crush food items and move them towards the mouth. The posterior five thoracic appendages were used for swimming, with the flap-like filaments undulating through the water column to create propulsion. These filaments were also likely used for breathing, like gills.

The predatory nature of Sidneyia is indicated by its spiny coxa used to masticate food, and the presence of crushed fossil debris in its gut. Sidneyia would have walked or swam above the sea floor, using its eyes and antennae to seek out prey, which it would capture and crush with its anterior appendages.

References:

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

BRIGGS, D. E. G. AND R. A. ROBISON. 1984. Exceptionally preserved non-trilobite arthropods and Anomalocaris from the Middle Cambrian of Utah. The University of Kansas Paleontological Contributions, 111: 1-24.

BRIGGS, D. E. G., B. S. LIEBERMAN, J. R. HENDRICKS, S. L. HALGEDAHL AND R. D. JARRARD. 2008. Middle Cambrian arthropods from Utah. Journal of Paleontology, 82(2): 238-254.

BRUTON, D. L. 1981. The arthropod Sidneyia inexpectans, Middle Cambrian, Burgess Shale, British Columbia. Philosophical Transactions of the Royal Society of London B, 295: 619-653.

BUDD, G. E. AND M. J. TELFORD. 2009. The origin and evolution of arthropods. Nature, 457(7231): 812-817.

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

CARON, J.-B., R. GAINES, G. MANGANO, M. STRENG, AND A. DALEY. 2010. A new Burgess Shale-type assemblage from the “thin” Stephen Formation of the Southern Canadian Rockies. Geology, 38: 811-814.

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.

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. Phylogenetic insights into the Cambrian radiation of arachnomorph arthropods. Journal of Paleontology, 82: 585-594.

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

RASSER, C. E. AND B. F. HOWELL. 1938. Lower Cambrian Olenellus zone of the Appalachians. Bulletin of the Geological Society of America, 49: 195-248.

SIMONETTA, A. M. 1963. Osservazioni sugli artropodi non trilobiti della Burgess Shale (Cambriano medio). II. Contributo: I Generai Sidneyia ed Amiella Walcott 1911. Monitore Zoologico Italiano, 70: 97-108.

STØMER, L. 1944. On the relationships and phylogeny of fossil and recent Arachnomorpha. Norsk Videnskaps-Akademi Skrifter I. Matematisk-Naturvidenskaplig Klasse, 5: 1-158.

WALCOTT, C. D. 1911. Middle Cambrian Merostomata. Cambrian geology and paleontology II. Smithsonian Miscellaneous Collections, 57: 17-40.

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

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, pp. 33-105. In G. D. Edgecombe (ed.), Arthropod fossils and phylogeny. Columbia University Press, New York.

ZHU, X., H. JIAN AND S. DEGAN. 2002. New occurrence of the Burgess Shale arthropod Sidneyia in the Early Cambrian Chengjiang Lagerstätte (South China), and revision of the arthropod Urokodia. Alcheringa: An Australasian Journal of Palaeontology, 26: 1-18.

Other Links:

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



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Sarotrocercus oblita

Reconstruction of Sarotrocercus oblita.

© MARIANNE COLLINS

Taxonomy:

Kingdom: 2D Model
Phylum: 2D Model
Higher Taxonomic assignment: Unranked clade (stem group arthropods)
Species name: Sarotrocercus oblita
Remarks:

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

Described by: Whittington
Description date: 1981
Etymology:

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

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

Type Specimens: Holotype –USNM144890 (part) and UNSM 272171 (counterpart) in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.
Other species:

Burgess Shale and vicinity: none.

Other deposits: none.

Age & Localities:

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:

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

Description:

Morphology:

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

Abundance:

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

Maximum Size:
16 mm

Ecology:

Life habits: 2D Model
Feeding strategies: 2D Model
Ecological Interpretations:

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

References:

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

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

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

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

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

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

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

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

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

Other Links:

None



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Hurdia victoria

3D animation of Hurdia victoria.

Animation by Phlesch Bubble © Royal Ontario Museum

Taxonomy:

Kingdom: 2D Model
Phylum: 2D Model
Higher Taxonomic assignment: Radiodonta, Family Hurdiidae
Species name: Hurdia victoria
Remarks:

With its single pair of jointed frontal appendages, lateral swimming flaps, and circular mouth structure, Hurdia possesses all the hallmarks of Radiodonta, part of the stem group to the true arthropods which also includes the iconic Anomalocaris (Collins, 1996). It is the namesake of the radiodontan family Hurdiidae, characterized by frontal appendages with comb or rake-like inner spines (Lerosey-Aubril & Pates, 2018). Hurdia is part of a diverse subgroup of hurdiids with large, elaborate carapaces (Caron & Moysiuk, 2021).

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 the Langston and Wheeler Formations of Utah (Lerosey-Aubril et al., 2020; Pates et al., 2018), and possibly the Pioche Formation of Nevada (Pates, Daley, Edgecombe, et al., 2019), the Jince Formation in the Czech Republic (Chlupáč & Kordule, 2002). and the Shuijingtuo Formation in Hubei Province, China (Cui & Huo, 1990).

Age & Localities:

Age:
Middle Cambrian, Wuliuan Stage, Burgess Shale Formation (around 505 million years old).
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 and the Monarch.

History of Research:

Brief history of research:

Although isolated parts of the body of Hurdia 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 crustacean (Rolfe, 1962). Hurdia’s frontal appendages were first described by Walcott (1911b) 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 radiodontans, the mouthparts were first described as the jellyfish Peytoia nathorsti (Walcott, 1911a). Whittington and Briggs (1985) discovered the first whole body specimens of Peytoia, associating the mouthparts with the former “Appendage F”. When describing Peytoia and 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 radiodontan, which also had an “Appendage F” pair and a frontal carapace complex 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 four distinct morphologies, two of which belong to Hurdia (known from two species, H. victoria and H. triangulata), one to Peytoia nathorsti, and one to an as yet unnamed species (Daley et al., 2013; Daley & Budd, 2010).

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-biomineralized carapaces consisting of one dorsal triangular H-element (previously called Hurdia) and two lateral paddle-shaped P-elements (from Proboscicaris). These carapaces wrap around the anterior margin of the head with the H-element spine protruding 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 notches in the overlapping posterior corners of the H- and P-elements. Hurdia triangulata differs from Hurdia victoria by having a wider and shorter H-element (Daley et al., 2013). Mouthparts 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 central 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 outer spines and eight elongated ventral spines (Daley et al., 2013; Pates, Daley, & Butterfield, 2019). The posterior half of the body consists of a series of seven to nine segments that extend laterally into triangular flaps. Recently, Van Roy and colleagues (2015) suggested that a second row of flaps might also be present in Hurdia. Each body segment is associated with a band of elongated blades interpreted to be gill structures. The body terminates abruptly in two rounded lobes (Daley et al., 2013). Complete specimens are up to 20 cm in length, although disarticulated fragments suggest a maximum body size up to 30 cm long (Caron & Moysiuk, 2021).

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:
300 mm

Ecology:

Life habits: 2D Model
Feeding strategies: 2D Model
Ecological Interpretations:

Hurdia was likely an active swimmer. Like other radiodontans, the animal propelled itself through the water column by undulating its flaps (Usami, 2006) and respiration would have been accomplished through the multiple rows of gills (Daley et al., 2013). The large dorsal carapace, upward facing eyes, and stubby body suggest it spent most of its time near the sea floor and may have fed primarily on burrowing organisms (Moysiuk & Caron, 2019). The function of the frontal carapace remains unknown, although it has been speculated to have played a role in feeding (Caron & Moysiuk, 2021; Daley et al., 2013). Prey items were funneled towards the mouth by a sweeping motion of the of the frontal appendages, with their rake-like inner spines forming a rigid net or cage (de Vivo et al., 2021; Moysiuk & Caron, 2019). Compared to Cambroraster and Titanokorys, the widely spaced spines on Hurdia’s appendages suggest it fed on relatively larger prey items (Caron & Moysiuk, 2021).

References:

  • BRIGGS, D. E. G. (1979). Anomalocaris: The largest known Cambrian arthropod. Palaeontology, 22(3), 631–664.
  • CARON, J.-B., & MOYSIUK, J. (2021). A giant nektobenthic radiodont from the Burgess Shale and the significance of hurdiid carapace diversity. Royal Society Open Science, 8(9), 210664. https://doi.org/10.1098/RSOS.210664
  • CHLUPÁČ, I., & KORDULE, V. (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(2), 280–293. https://doi.org/10.1017/S0022336000023362
  • CUI, Z.-L., & HUO, S.-C. (1990). New discoveries of Lower Cambrian crustacean fossils from western Hubei. Acta Palaeontologica Sinica, 29, 321–330.
  • DALEY, A. C., & BUDD, G. E. (2010). New anomalocaridid appendages from the Burgess Shale, Canada. Palaeontology, 53(4), 721–738. https://doi.org/10.1111/j.1475-4983.2010.00955.x
  • DALEY, A. C., BUDD, G. E., & CARON, J.-B. (2013). Morphology and systematics of the anomalocaridid arthropod Hurdia from the Middle Cambrian of British Columbia and Utah. Journal of Systematic Palaeontology, 11(7), 743–787. https://doi.org/10.1080/14772019.2012.732723
  • DALEY, A. C., BUDD, G. E., CARON, J.-B., EDGECOMBE, G. D., & COLLINS, D. (2009). The Burgess Shale anomalocaridid Hurdia and its significance for early euarthropod evolution. Science, 323(5921), 1597–1600. https://doi.org/10.1126/science.1169514
  • DE VIVO, G., LAUTENSCHLAGER, S., & VINTHER, J. (2021). Three-dimensional modelling, disparity and ecology of the first Cambrian apex predators. Proceedings of the Royal Society B, 288(1955). https://doi.org/10.1098/RSPB.2021.1176
  • LEROSEY-AUBRIL, R., KIMMIG, J., PATES, S., SKABELUND, J., WEUG, A., & ORTEGA-HERNÁNDEZ, J. (2020). New exceptionally preserved panarthropods from the Drumian Wheeler Konservat-Lagerstätte of the House Range of Utah. Papers in Palaeontology, 6(4), 501–531. https://doi.org/10.1002/spp2.1307
  • LEROSEY-AUBRIL, R., & PATES, S. (2018). New suspension-feeding radiodont suggests evolution of microplanktivory in Cambrian macronekton. Nature Communications, 9(1), 3774. https://doi.org/10.1038/s41467-018-06229-7
  • MOYSIUK, J., & CARON, J.-B. (2019). A new hurdiid radiodont from the Burgess Shale evinces the exploitation of Cambrian infaunal food sources. Proceedings of the Royal Society B, 286(1908), 20191079. https://doi.org/10.1098/rspb.2019.1079
  • PATES, S., DALEY, A. C., & BUTTERFIELD, N. J. (2019). First report of paired ventral endites in a hurdiid radiodont. Zoological Letters, 5(1), 18. https://doi.org/10.1186/s40851-019-0132-4
  • PATES, S., DALEY, A. C., EDGECOMBE, G. D., CONG, P., & LIEBERMAN, B. S. (2019). Systematics, preservation and biogeography of radiodonts from the southern Great Basin, USA, during the upper Dyeran (Cambrian Series 2, Stage 4). Papers in Palaeontology, 7(1), 235–262. https://doi.org/10.1002/spp2.1277
  • PATES, S., DALEY, A. C., & LIEBERMAN, B. S. (2018). Hurdiid radiodontans from the middle Cambrian (Series 3) of Utah. Journal of Paleontology, 92(1), 99–113. https://doi.org/10.1017/jpa.2017.11
  • ROLFE, W. D. (1962). Two new arthropod carapaces from the Burgess Shale (Middle Cambrian) of Canada. Breviora Museum of Comparative Zoology, 60, 1–9.
  • USAMI, Y. (2006). Theoretical study on the body form and swimming pattern of Anomalocaris based on hydrodynamic simulation. Journal of Theoretical Biology, 238, 11–17.
  • VAN ROY, P., DALEY, A. C., & BRIGGS, D. E. G. (2015). Anomalocaridid trunk limb homology revealed by a giant filter-feeder with paired flaps. Nature, 522(7554), 77. https://doi.org/10.1038/nature14256
  • WALCOTT, C. D. (1911a). Middle Cambrian holothurian and medusae. Smithsonian Miscellaneous Collections, 57, 41–68.
  • WALCOTT, C. D. (1911b). Middle Cambrian Merostomata. Cambrian Geology and Paleontology II. Smithsonian Miscellaneous Collections, 57, 17–40.
  • WALCOTT, C. D. (1912). Middle Cambrian Branchiopoda, Malacostraca, Trilobita and Merostomata. Smithsonian Miscellaneous Collections, 57, 145–228.
  • WHITTINGTON, H. B., & BRIGGS, D. E. G. (1985). The largest Cambrian animal, Anomalocaris, Burgess Shale, British Columbia. Philosophical Transactions of the Royal Society B: Biological Sciences, 309(1141), 569–609. https://doi.org/10.1098/rstb.1985.0096
Other Links:

http://www.sciencemag.org/content/323/5921/1597.abstract



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Habelia? brevicauda

Habelia? brevicauda (USNM 144910) – Holotype. Complete individual preserved without appendages. Total specimen length = 50 mm. Specimen dry – polarized light. Walcott Quarry.

© Smithsonian Institution – National Museum of Natural History. Photo: Jean-Bernard Caron

Taxonomy:

Kingdom: 2D Model
Phylum: 2D Model
Higher Taxonomic assignment: Unranked clade (stem group arthropods)
Species name: Habelia? brevicauda
Affinity:

Habelia? brevicauda is too poorly known to definitively determine its affinities. It has been aligned in some studies with the arachnomorphs (a group including chelicerates and trilobites), and has been suggested to be closely related to lamellipedians such as Naraoia and the trilobites (Briggs and Fortey, 1989), or placed within Megacheira as a close relative of Leanchoilia (Wills et al., 1998).

Described by: Simonetta
Description date: 1964
Etymology:

Habelia – from Mount Habel (3,161 m), today known as Mount Des Poilus, at the head of Yoho Valley, named in 1900 by Norman Collie in honour of Jean Habel, a German mountaineer. The name Mount Habel is now applied to a peak north of Mount Des Poilus.

brevicauda – from the Latin brevis, “short,” and cauda, “tail.”

Type Specimens: Holotype –USNM144910 in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.
Other species:

Burgess Shale and vicinity: Habelia optata from Walcott Quarry, Fossil Ridge and The Monarch in Kootenay National Park.

Other deposits: none.

Age & Localities:

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

The Walcott and Raymond Quarries on Fossil Ridge.

History of Research:

Brief history of research:

Habelia optata was first described by Walcott in 1912, to which Simonetta added the possible second species Habelia? brevicauda in 1964. This second species was later restudied by Whittington (1981). Phylogenetic analyses suggest a position within the arachnomorphs (Briggs and Fortey, 1989; Wills et al., 1998). If this is confirmed, Habelia probably represent a stem group of the Mandibulata, which includes crustaceans, myriapods, and hexapods (Scholtz and Edgecombe, 2006).

Description:

Morphology:

The body ranges in size from 1.8 – 5.4 cm and consists of a half-circle head shield and a trunk with twelve segments, the last of which bears a posterior spine. The head shield is smooth and featureless. The trunk segments have a broad, convex axial region, with blade-shaped elements (pleura) extending from either side. The pleura are short and round at the anterior of the body, but become progressively wider and have increasingly backward-pointing tips towards the posterior. The short, broad posterior spine tapers with a bluntly rounded tip.

In the type species, Habelia optata, the exoskeleton is covered in small tubercules , and appendages include a pair of antennae, two pairs of head appendages that are segmented and branch into two (biramous), and six pairs of possibly gnathobasic biramous trunk appendages (i.e., with a robust and spiny basal podomere or segment used for crushing food items). Tubercules and appendages have not been described in Habelia? brevicauda, which is why its placement in the genus is uncertain.

Abundance:

Habelia? brevicauda was originally described from fewer than ten specimens.

Maximum Size:
54 mm

Ecology:

Life habits: 2D Model
Feeding strategies: 2D Model
Ecological Interpretations:

Habelia? brevicauda is assumed to have walked on trunk limbs, using its head appendages to manipulate food items. If gnathobases were present, they may have served to masticate food. The frontal antennae were presumably sensory. Considerable flexure of the head may have been possible, which may have allowed Habelia to use its cephalon to dig into the sediment in search of food. It walked along the sea floor while digging and scavenging food items.

References:

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

ELLIOTT, D. K. AND D. L. MARTIN. 1987. A new trace fossil from the Cambrian Bright Angel Shale, Grand Canyon, Arizona. Journal of Paleontology, 61: 641-648.

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.

SIMONETTA, A. M. 1964. Osservazioni sugli artropodi non trilobiti della ‘Burgess Shale’ (Cambriano medio). III conributo. Monitore Zoologico Italiano, 72: 215-231.

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

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.

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

Other Links:

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Habelia optata

Reconstruction of Habelia optata.

© Marianne Collins

Taxonomy:

Kingdom: 2D Model
Phylum: 2D Model
Higher Taxonomic assignment: Unranked clade (stem group arthropods)
Species name: Habelia optata
Remarks:

Habelia optata is an arthropod, but its exact relationships remain poorly understood. It has been aligned in some studies to the arachnomorphs (a group including chelicerates and trilobites), and has either been allied with lamellipedians such as Naraoia and the trilobites (Briggs and Fortey, 1989), or placed within Megacheira as closely related to Leanchoilia (Wills et al., 1998).

Described by: Walcott
Description date: 1912
Etymology:

Habelia – from Mount Habel (3,161 m), today known as Mount Des Poilus, at the head of Yoho Valley. Named in 1900 by Norman Collie in honour of Jean Habel, a German mountaineer. The name Mount Habel is now applied to a peak north of Mount Des Poilus.

optata – unspecified; may derive from the Latin optatus, “wish or desire.”

Type Specimens: Holotype –USNM57693 in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.
Other species:

Burgess Shale and vicinity: Habelia? brevicauda from Walcott Quarry and Raymond Quarry, Fossil Ridge.

Other deposits: none.

Age & Localities:

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

The Walcott and Raymond Quarries on Fossil Ridge.

History of Research:

Brief history of research:

Habelia optata was first described by Walcott in 1912, and a possible second species Habelia? brevicauda was added to the genus by Simonetta in 1964. Habelia was later restudied by Whittington (1981). Habelia has been included in some phylogenetic analyses of arthropod relationships (Briggs and Fortey, 1989; Wills et al., 1998) and unusual zig-zag fossil tracks from the Middle Cambrian of the Grand Canyon have been ascribed to an arthropod similar to Habelia (Elliott and Martin, 1987).

Description:

Morphology:

Habelia optata is unusual in that its entire body is covered in tubercles (small, rounded nodules) that are particularly dense on the head shield and the axis of the body trunk. Its body consists of a convex head shield without eyes, and twelve body tergites with a long, jointed posterior spine projecting from the twelfth segment. The first three tergites have a thick median spine that bore tubercles. The head has a pair of multi-segmented setose antennae at the front, and two pairs of possibly biramous appendages with segmented walking limbs and dark sheets that may be filamentous branches.

The twelve body segments have a thick, blunt median spine on the dorsal surface. The first six body segments have appendages that are segmented and branch into two (biramous), including long stout segmented gnathobasic walking limbs (i.e., with a robust and spiny basal podomere or segment used for crushing food items) and a lobed outer branch with lamellae (small elongated structures) along the margin. The lobes are also present on the posterior segments, but no walking branches are associated with them. The tail is a long spine with a single joint midway along its length.

Abundance:

Extremely rare

Maximum Size:
41 mm

Ecology:

Life habits: 2D Model
Feeding strategies: 2D Model
Ecological Interpretations:

Habelia optata probably used its six trunk limbs for walking, reserving the head appendages for manipulating food items. It is likely that the frontal antennae were used to sense the environment since there are no obvious eyes. The size and shape of the posterior margin of the head suggests that there was considerable flexure possible between the head and the body, indicating that Habelia may have dug in the sediment for food items. It lived on the muddy seafloor and was heavily protected against predators by its thick body armor and pointed posterior spine, the latter of which would make it difficult for predators to attack from behind.

References:

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

ELLIOTT, D. K. AND D. L. MARTIN. 1987. A new trace fossil from the Cambrian Bright Angel Shale, Grand Canyon, Arizona. Journal of Paleontology, 61: 641-648.

SIMONETTA, A. M. 1964. Osservazioni sugli artropodi non trilobiti della ‘Burgess Shale’ (Cambriano medio). III conributo. Monitore Zoologico Italiano, 72: 215-231.

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

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.

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

Other Links:

None



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Pikaia gracilens

3D animation of Pikaia gracilens.

ANIMATION BY PHLESCH BUBBLE © ROYAL ONTARIO MUSEUM

Taxonomy:

Kingdom: 2D Model
Phylum: 2D Model
Higher Taxonomic assignment: Unranked clade (stem group chordates)
Species name: Pikaia gracilens
Remarks:

Pikaia is considered to represent a primitive chordate (Conway Morris, 1979; Conway Morris et al., 1982) possibly close to craniates (Janvier, 1998); a stem-chordate (Smith et al., 2001); or a cephalochordate (Shu et al., 1999). Its exact position within the chordates is still uncertain and this animal awaits a full redescription.

Described by: Walcott
Description date: 1911
Etymology:

Pikaia – from the pika, a small alpine mammal and cousin of the rabbits. Pikas live in the Rocky Mountains, including near the Burgess Shale.

gracilens – from the Latin gracilens, “thin, simple,” in reference to the shape of the body.

Type Specimens: Syntypes –USNM57628b, 57629 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:

Pikaia was first described by Walcott based on a couple of specimens in a 1911 monograph dealing with various Burgess Shale worms. Two additional specimens were figured in a posthumous publication (Walcott, 1931). Walcott placed Pikaia in a now defunct group called the Gephyrea with other vermiform fossils such as Banffia, Ottoia and Oesia. Pikaia was later considered to be a primitive chordate (Conway Morris, 1979; Conway Morris et al., 1982), an interpretation which has since been followed to some degree in most discussions about early chordate evolution (e.g., Janvier, 1998). Pikaia played a major part in Gould’s interpretations of the Burgess Shale fossils in Wonderful Life (Gould, 1989; see also Briggs and Fortey, 2005). A full redescription of this animal is currently under way (Conway Morris and Caron, in prep.).

Description:

Morphology:

Pikaia resembles Metaspriggina in outline, another chordate animal from the Burgess Shale, with an elongate body and a small anterior region bearing the head. The body is laterally flattened and there is evidence of a ventral fin towards the posterior. Numerous V-shaped or ziz-zag segments interpreted as myomeres or muscle bands are visible in all specimens. A narrow dorsal structure which runs down the length of the organism might represent a notochord, but this interpretation remains to be confirmed. The head bears two equal lobes and a pair of short and slender tentacle-like structures. There is no evidence of eyes. Just behind the head, on the ventral side of the body, there is a series of up to twelve pairs of small, short, pointed structures on either side of the midline. These are thought to be related to gill openings. The gut is narrow and the anus is terminal.

Abundance:

Pikaia is relatively rare, known from more than 60 specimens, all from the Walcott Quarry where it represents 0.03% of the specimens counted in the community (Caron and Jackson, 2008).

Maximum Size:
55 mm

Ecology:

Life habits: 2D Model
Feeding strategies: 2D Model
Ecological Interpretations:

The eel-like morphology and musculature of the animal suggest that it was likely free-swimming, although it probably spent time on the sea floor. The tentacles may have had a sensory function, and the presence of mud in its gut suggests that Pikaia was potentially a deposit feeder.

References:

BRIGGS, D. E. G. AND R. A. FORTEY. 2005. Wonderful strife: Systematics, stem groups, and the phylogenetic signal of the Cambrian radiation. Paleobiology, 31(SUPPL.2 ): 94-112.

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

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. H. B. WHITTINGTON, D. E. G. BRIGGS, C. P. HUGHES AND D. L. BRUTON. 1982. Atlas of the Burgess Shale. Palaeontological Association, 31 p. + 23 pl.

GOULD, S. J. 1989. Wonderful Life. The Burgess Shale and the Nature of History. Norton, New York, 347 p.

JANVIER, P. 1998. Les vertébrés avant le Silurien. GeoBios, 30: 931-950.

SHU, D.-G,. H. L. LUO, S. CONWAY MORRIS, X. L. ZHANG, S. X. HU, L. CHEN, J. HAN, M. ZHU, Y. LI AND L. Z. CHEN. 1999. Lower Cambrian vertebrates from south China. Nature, 402(4 November 1999): 42-46.

SMITH, M. P., I. J. SANSOM AND K. D. COCHRANE. 2001. The Cambrian origin of vertebrates, p. 67-84. In P. E. Ahlberg (ed.), Major Events in Early Vertebrate Evolution: Palaeontology, Phylogeny, Genetics and Development. Taylor and Francis, London.

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://paleobiology.si.edu/burgess/pikaia.html