Anomalocaris

Anomalocaris
Temporal range: Early to mid Cambrian: Chengjiang–Burgess shale
Image of the first complete Anomalocaris fossil found, residing in the Royal Ontario Museum
Scientific classification
Kingdom:Animalia
Clade:Euarthropoda
Class:†Dinocaridida
Order:†Radiodonta
Family:†Anomalocarididae
Genus:†Anomalocaris
Whiteaves 1892
Species
  • ?A. briggsi Nedin, 1995
  • A. canadensis Whiteaves, 1892
  • A. pennsylvanica Resser, 1929 (= A. canadensis?)[1]
  • A. saron Hou, Bergström & Ahlberg, 1995
Synonyms

Defunct species:[1]

  • A. lineata Resser & Howell, 1938
  • A. nathorsti Walcott, 1911

Anomalocaris ("abnormal shrimp") is an extinct genus of anomalocaridid, a family of animals thought to be closely related to ancestral arthropods. The first fossils of Anomalocaris were discovered in the Ogygopsis Shale by Joseph Frederick Whiteaves, with more examples found by Charles Doolittle Walcott in the Burgess Shale.[2] Originally several fossilized parts discovered separately (the mouth, feeding appendages and tail) were thought to be three separate creatures, a misapprehension corrected by Harry B. Whittington and Derek Briggs in a 1985 journal article.[2][3]

Discovery

"Arm" (top) and mouthpiece (bottom) from British Columbia

Anomalocaris has been misidentified several times, in part due to its makeup of a mixture of mineralized and unmineralized body parts; the mouth and feeding appendage was considerably harder and more easily fossilized than the delicate body.[4] Its name originates from a description of a detached 'arm', described by Joseph Frederick Whiteaves in 1892, as a separate crustacean-like creature due to its resemblance to the tail of a lobster or shrimp.[4] The first fossilized anomalocaridid mouth was discovered by Charles Doolittle Walcott, who mistook it for a jellyfish and placed it in the genus Peytoia. Walcott also discovered a second feeding appendage, but failed to realize the similarities to Whiteaves' discovery and instead identified it as feeding appendage or tail of the extinct Sidneyia.[4]

The body was discovered separately and classified as a sponge in the genus Laggania; a mouth was found with the body, but was interpreted by its discoverer Simon Conway Morris as an unrelated Peytoia that had through happenstance settled and been preserved with Laggania. Later, while clearing what he thought was an unrelated specimen, Harry B. Whittington removed a layer of covering stone to discover the unequivocally connected arm thought to be a shrimp tail and mouth thought to be a jellyfish.[4][2] Whittington linked the two species, but it took several more years for researchers to realize that the continuously juxtaposed Peytoia, Laggania and feeding appendage actually represented a single, enormous creature. The two genera are now placed into the family Anomalocaridae.[4] Because Peytoia was named first, it became the correct name for the entire animal. The original feeding arm, however, came from a larger species distinct from Peytoia and "Laggania", which retains the name Anomalocaris.[5]

In 2011, six fossils of compound eyes, the first for Anomalocaris, were recovered from an paleontological dig at Emu Bay on Kangaroo Island, Australia, proving that Anomalocaris was indeed an arthropod as had been suspected. The find also indicated that advanced arthropod eyes had evolved very early, before the evolution of jointed legs or hardened exoskeletons.[6]

Stephen Jay Gould cites Anomalocaris as one of the fossilized extinct species he believed to be evidence of a much more diverse set of phyla that existed in the Cambrian Period,[4] discussed in his book Wonderful Life, a conclusion disputed by other paleontologists.[2]

Anatomy

Size comparison of Anomalocaris (red), a human, and contemporary animals

For the time in which it lived, Anomalocaris was gigantic, up to 1 metre (3.3 feet) long.[2] It propelled itself through the water by undulating the flexible lobes on the sides of its body.[7] Each lobe sloped below the one more posterior to it,[8] and this overlapping allowed the lobes on each side of the body to act as a single "fin", maximizing the swimming efficiency.[7] The construction of a remote-controlled model showed this mode of swimming to be intrinsically stable,[9] implying that Anomalocaris would not have needed a complex brain to manage balance while swimming. The body was widest between the third and fifth lobe and narrowed towards the tail; it had at least 11 lobes in total.[8] It is difficult to distinguish lobes near the tail, making an accurate count difficult.[8]

Anomalocaris had an unusual disk-like mouth. The mouth was composed of 32 overlapping plates, four large and 28 small, resembling a pineapple ring with the center replaced by a series of serrated prongs.[2] The mouth could constrict to crush prey, but never completely close, and the tooth-like prongs continued down the walls of the gullet.[4] Two large 'arms' (up to 18 centimetres (7.1 inches) in length when extended[4]) with barb-like spikes were in front of the mouth.[3] The tail was large and fan-shaped, and along with undulations of the lobes, was probably used to propel the creature through Cambrian waters.[2][7][10] Stacked lamella of what were probably gills attached to the top of each lobe.

The eyes were 30 times as powerful as those of trilobites, long thought to have had the most advanced eyes of any contemporary species. With 16,000 lenses, the resolution of the 3-centimetre-wide (1.2 in) eyes would have been rivalled only by that of the modern dragonfly, which has 28,000 lenses in each eye.[6]

Paleobiology

Diet

Reconstruction of A. canadensis

A long-standing view holds that Anomalocaris fed on hard-bodied animals, including trilobites–making it one of the first predators–and its mid-gut glands strongly suggest a predatory lifestyle. However, its ability to penetrate mineralised shells has been questioned.[11] Some Cambrian trilobites have been found with round or W-shaped "bite" marks, which were identified as being the same shape as the mouthparts of Anomalocaris.[12] Stronger evidence that Anomalocaris ate trilobites comes from coprolite, which contain trilobite parts and are so large that the anomalocarids are the only known organism from that period large enough to have produced them.[12] However, since Anomalocaris lacks any mineralised tissue, it seemed unlikely that it would be able to penetrate the hard, calcified shell of trilobites.[12] Rather, the coprolites may have been produced by a different organism, such as a trilobite of the genus Redlichia.[13]

The lack of wear on anomalocaridid mouthparts suggests they did not come into regular contact with mineralised trilobite shells, and were possibly better suited to feeding on smaller, soft-bodied organisms by suction, since they would have experienced structural failure if they were used against the armour of trilobites.[11][13] A. canadensis may have been capable of feeding on organisms with hard exoskeletons due to the short, robust spines on its appendages.[13] However, as opposed to Peytoia whose mouthpiece is more rectangular with short protruding spines, the mouthpiece of A. canadensis has a smaller and more irregular diamond-shaped opening, not permitting strong biting motions, and indicating a suction-feeding behavior to suck in softer organisms.[14]

Anomalocaris model, National Dinosaur Museum, Canberra, Australia.

Phylogenetic analyses have also found "A." briggsi to belong to the closely related, filter-feeding family Cetiocaridae, which suggests that may have been a filter-feeder as well.[15][16]

One suggested possibility is that anomalocaridids fed by grabbing one end of their prey in their jaws while using their appendages to quickly rock the other end of the animal back and forth. This produced stresses that exploited the weaknesses of arthropod cuticles, causing the prey's exoskeleton to rupture and allowing the predator to access its innards.[12] This behaviour was originally thought to have provided an evolutionary pressure for trilobites to roll up, to avoid being flexed until they snapped.[12]

Paleoecology

Anomalocaris had a cosmopolitan distribution in Cambrian seas, and has been found from early to mid Cambrian deposits of Canada, the United States, China, and Australia.[17][18][19]

See also

Footnotes

  1. 1 2 Daley, A. C.; Peel, J. S. (2010). "A Possible Anomalocaridid from the Cambrian Sirius Passet Lagerstätte, North Greenland". Journal of Paleontology. 84 (2): 352–355. doi:10.1666/09-136R1.1.
  2. 1 2 3 4 5 6 7 Conway Morris, S. (1998). The crucible of creation: the Burgess Shale and the rise of animals. Oxford [Oxfordshire]: Oxford University Press. pp. 56–9. ISBN 0-19-850256-7.
  3. 1 2 Whittington, H.B.; Briggs, D.E.G. (1985). "The largest Cambrian animal, Anomalocaris, Burgess Shale, British Columbia". Philosophical Transactions of the Royal Society B. 309 (1141): 569–609. Bibcode:1985RSPTB.309..569W. doi:10.1098/rstb.1985.0096.
  4. 1 2 3 4 5 6 7 8 Gould, Stephen Jay (1989). Wonderful life: the Burgess Shale and the nature of history. New York: W.W. Norton. pp. 194–206. ISBN 0-393-02705-8.
  5. ↑ Daley, A. and Bergström, J. (2012). "The oral cone of Anomalocaris is not a classic 'peytoia'." Naturwissenschaften, doi:10.1007/s00114-012-0910-8
  6. 1 2 Paterson, J. R.; García-Bellido, D. C.; Lee, M. S. Y.; Brock, G. A.; Jago, J. B.; Edgecombe, G. D. (2011). "Acute vision in the giant Cambrian predator Anomalocaris and the origin of compound eyes". Nature. 480 (7376): 237–240. Bibcode:2011Natur.480..237P. doi:10.1038/nature10689. PMID 22158247.
  7. 1 2 3 Usami, Yoshiyuki (2006). "Theoretical study on the body form and swimming pattern of Anomalocaris based on hydrodynamic simulation". Journal of Theoretical Biology. 238 (1): 11–17. doi:10.1016/j.jtbi.2005.05.008. PMID 16002096.
  8. 1 2 3 Whittington, H.B.; Briggs, D.E.G. (1985). "The Largest Cambrian Animal, Anomalocaris, Burgess Shale, British Columbia". Philosophical Transactions of the Royal Society B (free full text). 309 (1141): 569–609. Bibcode:1985RSPTB.309..569W. doi:10.1098/rstb.1985.0096.
  9. ↑ Briggs, Derek E. G. (1994). "Giant Predators from the Cambrian of China". Science. 264 (5163): 1283–4. Bibcode:1994Sci...264.1283B. doi:10.1126/science.264.5163.1283. PMID 17780843.
  10. ↑ "The Anomalocaris homepage". Archived from the original on 4 April 2008. Retrieved 2008-03-20.
  11. 1 2 Hagadorn, James W. (August 2009). "Taking a Bite out of Anomalocaris" (PDF). In Smith, Martin R.; O'Brien, Lorna J.; Caron, Jean-Bernard. Abstract Volume. International Conference on the Cambrian Explosion (Walcott 2009). Toronto, Ontario, Canada: The Burgess Shale Consortium (published 31 July 2009). ISBN 978-0-9812885-1-2.
  12. 1 2 3 4 5 Nedin, C. (1999). "Anomalocaris predation on nonmineralized and mineralized trilobites". Geology. 27 (11): 987–990. Bibcode:1999Geo....27..987N. doi:10.1130/0091-7613(1999)027<0987:APONAM>2.3.CO;2.
  13. 1 2 3 Daley, A. C.; Paterson, J. R.; Edgecombe, G. D.; García-Bellido, D. C.; Jago, J. B. (2013). Donoghue, Philip, ed. "New anatomical information on Anomalocaris from the Cambrian Emu Bay Shale of South Australia and a reassessment of its inferred predatory habits". Palaeontology. doi:10.1111/pala.12029.
  14. ↑ Daley, A. C.; Bergström, J. (2012). "The oral cone of Anomalocaris is not a classic peytoia". Naturwissenschaften. 99 (6): 501–504. Bibcode:2012NW.....99..501D. doi:10.1007/s00114-012-0910-8. PMID 22476406.
  15. ↑ Vinther, J.; Stein, M.; Longrich, N. R.; Harper, D. A. T. (2014). "A suspension-feeding anomalocarid from the Early Cambrian". Nature. 507: 496–499. Bibcode:2014Natur.507..496V. doi:10.1038/nature13010. PMID 24670770.
  16. ↑ 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: 77–80. Bibcode:2015Natur.522...77V. doi:10.1038/nature14256. PMID 25762145.
  17. ↑ Anomalocaris at fossilworks.org (retrieved 30 September 2018)
  18. ↑ Briggs, D. E. G.; Mount, J. D. (1982). "The Occurrence of the Giant Arthropod Anomalocaris in the Lower Cambrian of Southern California, and the Overall Distribution of the Genus". Journal of Paleontology. 56 (5): 1112–8. JSTOR 1304568.
  19. ↑ Briggs, D. E. G.; Robison, R. A. (1984). "Exceptionally preserved nontrilobite arthropods and Anomalocaris from the Middle Cambrian of Utah". University of Kansas Paleontological Contributions (111). ISSN 0075-5052.
  • "Anomalocaris canadensis". Burgess Shale Fossil Gallery. Virtual Museum of Canada. 2011.
  • Anomalocaris 'homepage' with swimming animation
  • Burgess Shale: Anomalocaris canadensis (proto-arthropod), Smithsonian.
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