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Kootenia burgessensis

Trilobite with blunt head spines and smooth tail

Kootenia burgessensis (ROM 60761). Disarticulated specimen. Specimen dry – direct light (left) and coated with ammonium chloride sublimate to show details (right). Specimen length = 44 mm. Walcott Quarry.

© Royal Ontario Museum. Photo: Jean-Bernard Caron

Taxonomy:

Kingdom: Animalia
Phylum: Arthropoda
Higher Taxonomic assignment: Class Trilobita, Order Corynexochida
Species name: Kootenia burgessensis
Remarks:

Trilobites are, by a large margin, the most diverse of extinct arthropod classes, encompassing more than 2000 species, although the biomineralization of their cuticle is likely largely responsible for their much better-known diversity. A recent probabilistic analysis of trilobite relationships suggests that many of the diagnostic traits used for the taxonomy of the main groups remain highly variable across lineages (Paterson, Edgecombe & Lee, 2019), which explains long-standing difficulties in resolving the trilobite evolutionary tree. The kinship of trilobites relative to other arthropod groups has been a historical dilemma also (Cotton & Braddy, 2004; Aria, Caron & Gaines, 2015; Paterson, 2020). Antennules and some other characters, on the one hand, would ally them with mandibulate arthropods (the Gnathomorpha or Antennulata hypothesis), while others would place them within the chelicerate lineage (the Arachnomorpha hypothesis). Recent phylogenetic results have brought increased support to Arachnomorpha, but there is still no consensus on the question (Aria, 2022).

Described by: Resser
Description date: 1942
Etymology:

Burgess Shale and vicinity: Kootenia dawsoni. However, see below regarding a possible synonymy with the genus Olenoides.

Other deposits: other species attributed to Kootenia are widespread in the Cambrian of North America, and have been recorded in Greenland, China, Australia, and elsewhere.

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: Kootenia dawsoni; Olenoides serratus. (Species of Kootenia are no longer considered different enough from those in Olenoides to warrant placement in a separate genus, but Kootenia is retained here for ease of reference to historical literature).

Other deposits: other species attributed to Kootenia are widespread in the Cambrian of North America, and have been recorded in Greenland, China, Australia, and elsewhere.

Age & Localities:

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

The Walcott Quarry on Fossil Ridge, and nearby localities on Mount Field; K. dawsoni is known from the Trilobite Beds and elsewhere on Mount Stephen.

History of Research:

Brief history of research:

Kootenia burgessensis was established by Charles Resser based on material Walcott included in K. dawsoni. Kootenia originally appeared as a subgenus of Bathyuriscus in Walcott’s 1889 paper revising many of Rominger’s Mount Stephen trilobite identifications. Walcott named B. (Kootenia) dawsoni after G. M. Dawson of the Geological Survey of Canada as a replacement for what Rominger had illustrated as Bathyurus (?) in 1887. In 1908, Walcott followed G. F. Matthew (1899) in calling this Dorypyge (Kootenia) dawsoni, but regarded Kootenia as a full genus in 1918. Harry Whittington included Kootenia burgessensis in his 1975 redescription of Burgess Shale appendage-bearing trilobites, illustrating a single specimen showing biramous thoracic limbs on one side. In 1994, Melzak and Westrop pointed out that the diagnostic character of Kootenia (depth of “interpleural” furrows on the pygidium) showed intraspecific variability, and argued that Kootenia—and perhaps other dorypygids trilobites—may have to be subsumed within Olenoides. We maintain Kootenia as a separate genus here pending formal taxonomic clarification.

Description:

Morphology:

Hard parts: adult dorsal exoskeletons may reach 5.5 cm in length and are broadly oval in outline. In most general features, Kootenia burgessensis resembles the co-occurring Olenoides serratus, with a semi-circular cephalon bearing genal spines, a thorax of seven segments, and a semi-circular pygidium. In Kootenia, however, spines on the thoracic pleural tips and shorter and blunter, as are those around the margin of the pygidium; interpleural furrows on the pygidium are absent to very faint.

Unmineralized anatomy: based on evidence from just a few specimens, Kootenia burgessensis, like Olenoides serratus, had a pair of flexible, multi-jointed “antennae” followed by three pairs of biramous limbs on the cephalon. Pairs of similar biramous appendages were attached under each thoracic segment, with a smaller number under the pygidium. No specimens, however, show any evidence of posterior antenna-like cerci as in Olenoides.

Abundance:

Kootenia burgessensis is moderately common in the Walcott Quarry section on Fossil Ridge, as is Kootenia dawsoni in the Mount Stephen Trilobite Beds.

Maximum Size:
55 mm

Ecology:

Life habits: Benthic, Mobile
Feeding strategies: Deposit feeder, Scavenger
Ecological Interpretations:

Adult Kootenia burgessensis walked along the sea bed, possibly digging shallow furrows to locate small soft-bodied and weakly-shelled animals or carcasses. Kootenia could probably swim just above the sea bed for short distances. Tiny larvae and early juveniles probably swam and drifted in the water column.

References:

  • ARIA, C. (2022) The origin and early evolution of arthropods. Biological Reviews 97, 1786–1809.
  • ARIA, C., CARON, J.-B. & GAINES, R. (2015) A large new leanchoiliid from the Burgess Shale and the influence of inapplicable states on stem arthropod phylogeny. Palaeontology 58, 629–660.
  • COTTON, T.J. & BRADDY, S.J. (2004) The phylogeny of arachnomorph arthropods and the origin of the Chelicerata. Transactions of the Royal Society of Edinburgh-Earth Sciences 94, 169–193.
  • PATERSON, J.R. (2020) The trouble with trilobites: classification, phylogeny and the cryptogenesis problem. Geological Magazine 157, 35–46.
  • PATERSON, J.R., EDGECOMBE, G.D. & LEE, M.S.Y. (2019) Trilobite evolutionary rates constrain the duration of the Cambrian explosion. Proceedings of the National Academy of Sciences of the United States of America 116, 4394–4399.
  • MATTHEW, G. F. 1899. Studies on Cambrian faunas, No. 3. Upper Cambrian Fauna of Mount Stephen, British Columbia: The trilobites and worms. Transactions of the Royal Society of Canada, Series 2, Vol. 5, Section IV:39-66.
  • MELZAK, A. AND S. R. WESTROP. 1994. Mid-Cambrian (Marjuman) trilobites from the Pika Formation, southern Canadian Rocky Mountains, Alberta. Canadian Journal of Earth Sciences, 31:969-985.
  • RASETTI, F. 1951. Middle Cambrian stratigraphy and faunas of the Canadian Rocky Mountains. Smithsonian Miscellaneous Collections, 116 (5): 1-277.
  • RESSER, C. E. 1942. Fifth contribution to nomenclature of Cambrian trilobites. Smithsonian Miscellaneous Collections, 101 (15): 1-58.
  • RESSER, C. E. 1942. Fifth contribution to nomenclature of Cambrian trilobites. Smithsonian Miscellaneous Collections, 101 (15): 1-58.
  • 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.
  • 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. 1889. Description of new genera and species of fossils from the Middle Cambrian. United States National Museum, Proceedings for 1888:441-446.
  • WALCOTT, C. D. 1908. Mount Stephen rocks and fossils. Canadian Alpine Journal, 1: 232-248.
  • WALCOTT, C. 1918. Cambrian Geology and Paleontology IV. Appendages of trilobites. Smithsonian Miscellaneous Collections, 67(4): 115-216.
  • WHITTINGTON, H. B. 1975. Trilobites with appendages from the Middle Cambrian, Burgess Shale, British Columbia. Fossils and Strata, No. 4: 97-136.
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