The Burgess Shale

Liangshanella burgessensis

3D animation of Liangshanella burgessensis.

ANIMATION BY PHLESCH BUBBLE © ROYAL ONTARIO MUSEUM

Taxonomy:

Kingdom: Very abundant
Phylum: Very abundant
Higher Taxonomic assignment: Unranked clade (Order: Bradoriida, stem group arthropods)
Species name: Liangshanella burgessensis
Remarks:

Liangshanella is a bradoriid belonging to the family Svealutidae (Siveter and Williams, 1997). The bradoriids were traditionally compared to other bivalved arthropods, such as Recent ostracods (e.g. Sylvester-Bradley, 1961) and Cambrian phosphatocopids (e.g. Maas et al., 2003). However, they are thought to be in the stem-lineage or in a sister group position relative to the Crustaceans (e.g. Hou et al., 1996; Shu et al., 1999; Hou et al., 2010).

Described by: Siveter and Williams
Description date: 1997
Etymology:

Liangshanella – from Liangshan, a region in South Shaanxi, China.

burgessensis – from the Burgess Shale. The name is derived from Mount Burgess (2,599 m), a mountain peak in Yoho National Park. Mount Burgess was named in 1886 by Otto Klotz, the Dominion topographical surveyor, after Alexander Burgess, a former Deputy Minister of the Department of the Interior.

Type Specimens: Holotype –USNM272083 in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.
Other species:

Burgess Shale and vicinity: none.

Other deposits: L. circumbolina from the Flinders Ranges in South Australia; L. liangshanensis, L. rotundata, L. orbicularis, L. yunnanensis and L. baensis from southern China; L. lubrica from the Tongying Formation in Hubei, China; L. sayutinae from the Trans-Baikal area in the Russian Far-East and Greenland; L. birkenmajeri from Antarctica. See references in Siveter and Williams (1997).

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:

Liangshanella liangshanensis is the type species of this genus and was first described by Huo (1956) from Lower Cambrian rocks of south China. Further species have since been described in China (Zhang, 1974; Li, 1975; Qian and Zhang, 1983; Zhang, 2007), Russia and Greenland (Melnikova, 1988), Australia (Topper et al., in press) and Antarctica (Wrona, 2009). Liangshanella burgessensis from the Burgess Shale was described by Siveter and Williams (1997), and the genus has been included in studies on the biogeography, evolution and affinity of the bradoriids (e.g. Shu and Chen, 1994; Williams et al., 2007).

Description:

Morphology:

Like all bradoriids, Liangshanella burgessensis has a small bivalved carapace with a straight dorsal hinge held together by a band of cuticle. The carapaces range in length from 0.66 mm – 4.25 mm and were soft and unmineralized. The bivalved carapace of L. burgessensis is sub-circular, with the anterior end being slightly narrower than the posterior end. There is a marginal ridge along the lateral surface of the valves. A centrally situated, sub-circular muscle scar composed of numerous small pits can be seen inside the valve. No evidence of soft parts has been found.

Abundance:

Liangshanella burgessensis is known from thousands of specimens and is the most common taxon in the Walcott Quarry (11.8% of the community, Caron and Jackson, 2008).

Maximum Size:
10 mm

Ecology:

Life habits: Very abundant
Feeding strategies: Very abundant
Ecological Interpretations:

Liangshanella was likely epibenthic, living on and within the first few metres of the soft muddy seafloor. Like other bradoriids, Liangshanella was probably a deposit feeder, and may have even been scavenging or predating on microscopic non-mineralized animals (Williams et al., 2007). Most specimens of Liangshanella found are empty carapaces, being left over from when the animal moulted its exoskeleton. Bradoriids are extremely common in Cambrian rocks, suggesting they played important roles in recycling nutrients in the seabed (Shu et al., 1999). They were also important food sources for larger animals, as indicated by their common presence in coprolites (e.g. Vannier and Chen, 2005).

References:

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

HOU, X., D. J. SIVETER, M. WILLIAMS, D. WALOSSEK AND J. BERGSTRÖM. 1996. Preserved appendages in the arthropod Kunmingella from the early Cambrian of China: its bearing on the systematic position of the Bradoriida and the fossil record of the Ostracoda. Philosophical Transactions of the Royal Society, B351: 1131-1145.

HOU, X., M. WILLIAMS, D.J. SIVETER, D.J. SIVETER, R.J. ALDRIDGE AND R.S. SANSOM. 2010. Soft-part anatomy of the Early Cambrian bivalve arthropods Kunyangella and Kunmingella: significance for the phylogenetic relationships of Bradoriida. Proceedings of the Royal Society, B277: 1835-1841.

HUO, S. 1956. Brief notes on lower Cambrian Archaeostraca from Shensi and Yunnan. Acta Palaeontologica Sinica, 4: 425-445.

LI, Y. 1975. Cambrian Ostracoda and other new descriptions from Sichuan, Yunnan and Shaanxi. Professional Papers of Stratigraphy and Palaeontology, 2: 37-72.

MAAS, A., D. WALOSZEK AND K.J. MÜLLER. 2003. Morphology, ontogeny and phylogeny of the Phosphatocopina (Crustacea) from the Upper Cambrian “Orsten” of Sweden. Fossils and Strata, 49: 1-238.

MELNIKOVA, L. M. 1988. Nekotoryye bradoriidy (Crustacea) iz botomskogo yarusa vostochnogo Zabaykal’ya. Paleontologicheskiy Zhurnal, 1: 114-117.

QIAN, Y. AND S. ZHANG. 1983. Small shelly fossils from the Xihaoping Member of the Tongying Formation in Fangxian County of Hubei Province and their stratigraphical significance. Acta Palaeontologica Sinica, 22: 82-94

SHU, D. AND L. CHEN. 1994. Cambrian palaeobiogeography of Bradoriida. Journal of Southeast Asian Earth Sciences, 9: 289-299.

SHU, D., J. VANNIER, H. LUO, L. CHEN, X. ZHANG AND S. HU. 1999. Anatomy and lifestyle of Kunmingella (Arthropoda, Bradoriida) from the Chengjiang fossil Lagerstätte (Lower Cambrian, Southwest China). Lethaia, 35: 279-298.

SIVETER, D.J. AND M. WILLIAMS. 1997. Cambrian Bradoriid and Phosphatocopid Arthropods of North America. Special Papers in Palaeontology, 57: 1-69.

SYLVESTER-BRADLEY, P. C. 1961. Archaeocopida, p. Q100-103. In R. C. Moore, and C. W. Pitrat (Eds.), Treatise on Invertebrate Paleontology Part Q, Arthropoda 3, Crustacea, Ostracoda. Geological Society of America and University of Kansas Press, Boulder, Colorado and Lawrence, Kansas.

Other Links:

None

Ogygopsis klotzi

Ogygopsis klotzi (figure 1) illustrated by Rominger (1887) as Ogygia klotzi.

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Taxonomy:

Kingdom: Very abundant
Phylum: Very abundant
Higher Taxonomic assignment: Trilobita (Order: Corynexochida)
Species name: Ogygopsis klotzi
Remarks:

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

Described by: Rominger
Description date: 1887
Etymology:

Ogygopsis – from Ogygia, in Greek mythology, the 7th daughter of Amphion and Niobe; Ogygia was first used as a trilobite genus name in 1817.

klotzi – after Otto Klotz, the Dominion topographical surveyor who provided the fossils for Rominger’s study and description.

Type Specimens: Holotype (K. burgessensis) – USNM65511 in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA (Resser, 1942); Type status under review – (K. dawsoni), University of Michigan Museum of Paleontology, Ann Arbor, Michigan, USA.
Other species:

Burgess Shale and vicinity: Ogygopsis spinulosa Rasetti, 1951, from the slightly older Cathedral Formation on Mount Stephen.

Other deposits: species of Ogygopsis have now been described from elsewhere in the Cambrian of North America, as well as in Greenland and Siberia.

Age & Localities:

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

The Trilobite Beds and smaller localities on Mount Stephen.

History of Research:

Brief history of research:

Ogygopsis klotzi was first described in the same 1887 publication as several other important Mount Stephen trilobites. Rominger named the largest and most abundant species after Klotz, placing it in the genus Ogygia. The following year, Charles Walcott questioned this assignment, and in 1889 proposed the new genus name Ogygopsis. For decades afterwards, Ogygopsis was thought to be unique to the Mount Stephen Trilobite Beds, where it is the most conspicuous fossil on the mountain slope. In fact, Walcott designated the occurrence as the “Ogygopsis shale” in 1908, and subsequently named the Burgess Shale as its geographic equivalent (although Ogygopsis itself has never been found on Fossil Ridge!)

Description:

Morphology:

Hard parts: adult dorsal exoskeletons may be up to 13 cm long and are elongate oval in outline, with a large crescentic cephalon, a thorax of eight segments ending pointed blade-like tips, and a large semi-circular pygidium without spines. The glabella is long and barrel-shaped, smoothly rounded in front, reaching almost to the anterior border. Thin eye ridges angle back from near the front of the glabella to long narrow eyes located opposite glabellar mid-length. Free cheeks extend back into straight, short genal spines. The large pygidium has 9 axial rings decreasing in size backwards, followed by a terminal piece; 8 or 9 pairs of pygidial ribs become progressively shorter and more backwardly directed. The whole exoskeleton is mostly smooth externally, with fine ridges parallel to the margins; free cheeks and posterior fixed cheeks may show a distinctive pattern of fine anastomosing (interlinking) ridges.

Unmineralized anatomy: only a very few specimens of Ogygopsis klotzi are known with preserved evidence of limbs, but these show that there was a pair of flexible, multi-jointed antennae on the cephalon (Hofmann and Parsley, 1966).

Abundance:

Ogygopsis klotzi is extraordinarily abundant at the Mount Stephen Trilobite Beds, where it is the most common fossil encountered (Rudkin, 1996; 2009), but it does not occur on Fossil Ridge. The vast majority of more-or-less complete specimens lack free cheeks, and many paleontologists have interpreted these as moulted individuals.

Maximum Size:
130 mm

Ecology:

Life habits: Very abundant
Feeding strategies: Very abundant
Ecological Interpretations:

The shape and size of Ogygopsis klotzi adults suggest that they walked along the sea bed. Because Ogygopsis occurs in such enormous numbers, it is hard to imagine it as a predator/scavenger, like Olenoides. It may instead have consumed much smaller organic particles in unusual environments. Obvious healed injuries have been described on a number of Ogygopsis specimens; some of these may be evidence of predation on freshly moulted “soft-shell” trilobites by larger arthropods such as Anomalocaris (Rudkin, 1979; 2009). The tiny larval stages and early juveniles of Ogygopsis probably swam and drifted in the water column above the sea bed.

References:

HOFFMAN, H. J. AND R. L. PARSLEY. 1966. Antennae of Ogygopsis. Journal of Paleontology, 40: 209-211.

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

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

RUDKIN, D. M. 1979. Healed injuries in Ogygopsis klotzi (Trilobita) from the Middle Cambrian of British Columbia. Royal Ontario Museum, Life Sciences Occasional Paper, 32: 1-8.

RUDKIN, D. M. 1996. The Trilobite Beds of Mount Stephen, Yoho National Park, p. 59-68. InR. Ludvigsen (ed.), Life in Stone – A Natural History of British Columbia’s Fossils. UBC Press, Vancouver.

RUDKIN, D. M. 2009. The Mount Stephen Trilobite Beds, p. 90-102. In J.-B. Caron and D. Rudkin (eds.), A Burgess Shale Primer – History, Geology, and Research Highlights. The Burgess Shale Consortium, Toronto.

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

WALCOTT, C. D. 1908. Mount Stephen rocks and fossils. Canadian Alpine Journal, 1: 232-248.

WHITTINGTON, H. B. 1975. Trilobites with appendages from the Middle Cambrian, Burgess Shale, British Columbia. Fossils and Strata, No. 4: 97-136.

Other Links:

http://www.trilobites.info/ordcorynexochida.htm

http://www.trilobites.info/trilovent.htm

Canadaspis perfecta

3D animation of Canadaspis perfecta.

Animation by Phlesch Bubble © Royal Ontario Museum

Taxonomy:

Kingdom: Very abundant
Phylum: Very abundant
Higher Taxonomic assignment: Unranked clade (stem group arthropods)
Species name: Canadaspis perfecta
Remarks:

Canadaspis was originally classified as a Malacostracan crustacean (Walcott, 1912; Briggs, 1978), but this has been widely debated (e.g. Hou and Bergström, 1997; Boxshall, 1998; Walossek, 1999; Butterfield, 2002). It has also been placed in the upper euarthropod stem-lineage (Edgecombe, 2010), forming a clade with other bivalved arthropods such as Perspicaris (Bergström and Hou, 1998; Waloszek et al., 2007), and possibly including Fuxianhuia (Budd, 2002; Budd and Telford, 2009).

Described by: Walcott
Description date: 1912
Etymology:

Canadaspis – from the country Canada, whose name derives from the Saint-Lawrence Iroquoian kanata, “settlement” or “land,” and the Greek aspis, “ shield.”

perfecta – from the Latin perfectus, “complete.”

Type Specimens: Lectotype –USNM57703 in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.
Other species:

Burgess Shale and vicinity: none.

Other deposits: C. laevigata from the Lower Cambrian Chengjiang biota (Hou and Bergström, 1991, 1997). Further material of Canadaspis cf. perfecta has been recovered from additional localities in the USA (Robison and Richards, 1981; Lieberman, 2003; Briggs et al., 2008).

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:

Originally referred to as Hymenocaris by Charles Walcott (1912), the genus Canadaspis was erected by Novozhilov (1960). Four species were designated by Simonetta and delle Cave (1975), but two of them, Canadaspis ovalis and Canadaspis dictynna have since been redescribed within Perspicaris dictynna (Briggs, 1977). A third species, Canadaspis obesa was redescribed within Canadaspis perfecta (Briggs, 1978). A full study of the fourth and only valid species, Canadaspis perfecta was published by Briggs (1978), who posited a crustacean affinity. This was rebuffed by later workers (Hou and Bergström, 1997; Boxshall, 1998; Walossek, 1999), and a close relationship with other bivalved Burgess Shale taxa in the arthropod stem lineage, including Branchiocaris, Perspicaris and Odaraia, was suggested (Budd, 2002; Budd, 2008).

Description:

Morphology:

Canadaspis is composed of a bivalved carapace covering a body with an appendage-bearing head region, an abdomen of 8 segments with associated limbs that are segmented and branch into two (biramous), and a thorax of 7 segments with a spiny telson or tail. The length of the bivalved carapaces ranges in size from 0.8-5.2 cm. The carapace valves are suboval in outline and taper towards the anterior, with a straight hinge line connecting them towards the back of the upper surface (dorsally).

The head has two pairs of antennae, small eyes, spiny mouth parts and two pairs of biramous appendages. The first antennae are short and unsegmented, while the second antennae are much longer, have at least 12 segments and are fringed with long spines. The small eyes were borne on short, blunt stalks. A series of spines behind the antennae are interpreted as mandibles, arthropod mouth parts used for cutting food. The ten pairs of biramous limbs of the head and abdomen consist of a segmented inner walking limb, and a large outer flap with lamellae, interpreted to be gills. The segmented abdomen does not bear appendages, and ends in a spiny telson. The gut of Canadaspis is sometimes preserved, with mid-gut glands giving it a segmented appearance (Butterfield, 2002).

Abundance:

Canadaspsis is abundant, with over 5,000 specimens known; it comprises 8.6% of the Walcott Quarry community (Caron and Jackson, 2008).

Maximum Size:
52 mm

Ecology:

Life habits: Very abundant
Feeding strategies: Very abundant
Ecological Interpretations:

Canadaspis was likely to have lived on the sea floor, walking on its biramous appendages by moving them in a rippling motion. This would also waft water past the gills that form the outer branches of its biramous limbs, allowing for respiration. This movement may also propelled Canadaspis through the water column. The biramous appendages on Canadapsis’ head are tipped with a pair of claws that were probably used in feeding. The inner surfaces of its legs were covered with spines that would have assisted in feeding by directing food particles to the organism’s mouth. The mandibles would have been used to help consume the coarse particles found on the sediment surface. Canadapsis’ spiny head-shield probably protected it from predators.

References:

BERGSTRÖM, J. AND X. HOU. 1998. Chengjiang arthropods and their bearing on early arthropod evolution, p. 151-184. In G. D. Edgecombe (ed.), Arthropod Fossils and Phylogeny. Columbia University Press, New York.

BOXSHALL, G. 1998. Comparative limb morphology in major crustacean groups: the coax-basis joint in postmandibular limbs, p. 155-167. In R. A Fortey and R. Thomas (eds.), Arthropod phylogeny. Chapman & Hall, London.

BRIGGS, D. E. G. 1977. Bivalved arthropods from the Cambrian Burgess Shale of British Columbia. Palaeontology, 20: 596-612.

BRIGGS, D. E. G. 1978. The morphology, mode of life, and affinities of Canadaspis perfecta (Crustacea: Phyllocarida), Middle Cambrian, Burgess Shale, British Columbia. Philosophical Transactions of the Royal Society of London, Series B, 281(984): 439-487.

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.

BUDD, G. E. 2002. A palaeontological solution to the arthropod head problem. Nature, 417(6886): 271-275.

BUDD, G. E. 2008. Head structure in upper stem-group euarthropods. Palaeontology, 51(3): 561-573.

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

BUTTERFIELD, N. J. 2002. Leanchoilia guts and the interpretation of three-dimensional structures in Burgess Shale-type fossils. Paleobiology, 28(1): 155-171.

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

EDGECOMBE, G. D. 2010. Arthropod phylogeny: An overview from the perspectives of morphology, molecular data and the fossil record. Arthropod Structure & Development, 39: 74-87.

HOU, X. AND J. BERGSTRÖM. 1991. The arthropods of the Lower Cambrian Chengjiang fauna, with relationships and evolutionary significance p. 179-187. In A. M. Simonetta and S. Conway Morris (eds.), The Early Evolution of Metazoa and the Significance of Problematic Taxa. Cambridge University Press, Cambridge.

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

LIEBERMAN, B. S. 2003. A new soft-bodied fauna: the Pioche Formation of Nevada. Journal of Paleontology, 77(4): 674-690.

NOVOZHILOV. N. I. 1960. Principles of Paleontology: arthropods, trilobites and crustaceans. In Y. A. Orlov (ed.). Gos. Nauchno-Techn. Izdvo, Moscow.

ROBISON, R. A. AND B. C. RICHARDS. 1981. Larger bivalve arthropods from the Middle Cambrian of Utah. The University of Kansas Paleontological Contributions, 106: 1-28.

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. D. 1912. Middle Cambrian Branchiopoda, Malacostraca, Trilobita, and Merostomata, pp. 145-228, Cambrian Geology and Paleontology, 2.Volume 57 (6). Smithsonian Miscellaneous Collections.

WALOSZEK, D. 1999. On the Cambrian diversity of Crustacea, p. 3-27. In F. R. Schram and J. C. von Vaupel Klein (eds.), Crustaceans and the biodiversity crisis. Volume 1. Brill, Leiden.

WALOSZEK, D. MAAS, A. CHEN, J. AND M. STEIN. 2007. Evolution of cephalic feeding structures and the phylogeny of Arthropoda. Palaeogeography, Palaeoclimatology, Palaeoecology, 254: 273-287.

Other Links:

Marrella splendens

3D animation of Marrella splendens.

ANIMATION BY PHLESCH BUBBLE © ROYAL ONTARIO MUSEUM

Taxonomy:

Kingdom: Very abundant
Phylum: Very abundant
Higher Taxonomic assignment: Marrellomorpha (Order: Marrellida, stem group arthropods)
Species name: Marrella splendens
Remarks:

The affinity of Marrella is still somewhat uncertain. It has been grouped together with the Devonian taxa Mimetaster and Vachonisia from the Hunsrück Shale to form the Class Marrellomorpha (Beurlen, 1934; Strømer, 1944), but the placement of this class in arthropod evolution is unclear. It has been suggested to be at the base of a group of Lamellipedian arthropods, including trilobites and trilobite-like taxa, (Hou and Bergström, 1997), but has also been placed in the most basal position in the upper stem lineage arthropods (Briggs and Fortey, 1989; Wills et al., 1998).

Described by: Walcott
Description date: 1912
Etymology:

Marrella – after Dr. John Marr, palaeontologist at Cambridge University and friend of Walcott.

splendens – from the Latin splendens, “beautiful, or brilliant.”

Type Specimens: Lectotype –USNM57674 in the National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.
Other species:

Burgess Shale and vicinity: none

Other deposits: Marrella sp. from the Kaili Biota of southwest China (Zhao et al., 2003).

Age & Localities:

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

The Walcott and Raymond Quarries on Fossil Ridge. Smaller localities on Mount Field, the Tulip Beds (S7) on Mount Stephen and Mount Odaray.

History of Research:

Brief history of research:

Marrella was one of the first fossils found by Walcott, and sketches appear in his notebook as early as August 31st, 1909. Walcott informally named them “lace crabs” at the time. The next summer, on August 9, 1910, Walcott and son Stuart found the “lace crab beds” in situ, marking the discovery of the fossil-bearing beds of the Walcott Quarry of the Burgess Shale. Walcott (1912) formally described the “lace crabs” as Marrella splendens, but a reconstruction was not attempted until Raymond (1920).

Marrella was examined again by Simonetta (1962) and in a major study by Whittington (1971). New specimens collected by the Royal Ontario Museum allowed for the description of a specimen showing Marrella in the act of moulting (García-Bellido and Collins, 2004), and another re-description of the taxon (García-Bellido and Collins, 2006).

Description:

Morphology:

Marrella is a small arthropod with a wedge-shaped head shield bearing two pairs of prominent spines that project from the sides and posterodorsal margin and extend back along most of the length of the body. There is also a pair of smaller posteroventral spines. The head bears a pair of long, thin antennae with as many as 30 segments, and a pair of paddle-like appendages with six segments and numerous bushy setae along the edges.

Behind the head, the body consists of 26 segments that are small and subcircular, each bearing a pair of biramous appendages. The walking branch of this appendage has six segments, and the second branch is made of tapering gills with long, slim filaments that attach near the base of the legs. The last twelve body segments have conspicuous internal projections that form a net below the body.

The tail is minute and pointed. The stomach is located in the head near the ventral mouth, and the intestine stretches most of the length of the body. Dark stains found around the body are suggested to be the gut contents that were squeezed out during preservation. A small, triangular dorsal heart is located in the cephalic region and has arteries branching off from it.

Abundance:

Marrella is one of the most common species in the Burgess Shale. Over 25,000 specimens have been collected (García-Bellido and Collins, 2006), and it is the second most common arthropod species in Walcott Quarry, comprising 7.3% of the specimens counted (Caron and Jackson, 2008).

Maximum Size:
25 mm

Ecology:

Life habits: Very abundant
Feeding strategies: Very abundant
Ecological Interpretations:

Marrella was an active swimmer that moved just above the sea floor while deposit feeding. It could rest on the sea floor by standing on its body appendages. Swimming was achieved by undulating the second pair of paddle-like appendages on the head. Its antennae would be used to sense the environment and locate food items. The net of internal projections on the last twelve body segments would have been used to trap food particles located in water currents and to pass them along the underside of the animal. Food particles trapped in the net would be moved towards the mouth using the tips of the anterior legs.

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., 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.

BEURLEN, K. 1934. Die Pygaspiden, eine neue Crustaceen – (Entomostraceen) – Gruppe aus den Mesosaurier führenden Iraty-Scichten Brasiliens. Paläontologische Zeitschrift, 16: 122-138.

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

GARCÍA-BELLIDO, D. AND D. H. COLLINS. 2004. Moulting arthropod caught in the act. Nature, 429: 40.

GARCÍA-BELLIDO, D. AND D. H. COLLINS. 2006. A new study of Marrella splendens(Arthropoda, Marrellomorpha) from the Middle Cambrian Burgess Shale, British Columbia, Canada. Canadian Journal of Earth Sciences, 43: 721-742.

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

RAYMOND, P. E. 1920. The appendages, anatomy, and relationships of trilobites. Memoirs of the Connecticut Academy of Arts and Sciences, 7: 1-169.

SIMONETTA, A. M. 1962. Note sugli artropodi non trilobiti della Burgess Shale, Cambriano Medio della Columbia Britannica (Canada). 1. contributo: 2. genere Marrella Walcott, 1912. Monitore Zoologico Italiano, 69: 172-185.

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. 1912. Cambrian geology and paleontology II. Middle Cambrian Branchiopoda, Malacostraca, Trilobita and Merostomata. Smithsonian Miscellaneous Collections, 57(6): 145-228.

WHITTINGTON, H. B. 1971. Redescription of Marrella splendens (Trilobitoidea) from the Burgess Shale, Middle Cambrian, British Columbia. Bulletin of the Geological Survey of Canada, 209: 1-24.

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.

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Other Links:

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