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Liangshanella burgessensis

A tiny, abundant arthropod with flat bivalved carapace

3D animation of Liangshanella burgessensis.

ANIMATION BY PHLESCH BUBBLE © ROYAL ONTARIO MUSEUM

Taxonomy:

Kingdom: Animalia
Phylum: Arthropoda
Higher Taxonomic assignment: Hymenocarines?, Order Bradoriida (Raymond 1935).
Species name: Liangshanella burgessensis
Remarks:

Liangshanella is a bradoriid belonging to the family Svealutidae (Siveter & Williams 1997). The bradoriids were traditionally compared to other bivalved arthropods within or close to crustaceans, such as Recent ostracods (e.g. Sylvester-Bradley, 1961) and Cambrian phosphatocopids (e.g. Maas et al. (2003)). More recently, authors have argued bradoriids to represent more basal members of the arthropod stem group (e.g. Hou et al. (1996); Shu et al. (1999); Hou et al. (2010)). A recent reexamination of appendage morphology in the bradoriid Kunmingella supports a placement of these arthropods outside of crown groups (Zhai et al. 2019), but their exact phylogenetic placement remains unclear.

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, Wuliuan stage, Burgess Shale Formation (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. urther 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). The most recent insight on the morphoanatomy of these arthropods comes from computed tomographic imagery of Kunmingella species from the Chengjiang biota (Zhai et al. 2019), which confirms both the presence of biramous enditic appendages and the lack of clear crown-group synapomorphies.

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. Based on appendicular evidence from other bradoriid taxa, L. burgessensis would have possessed short, spinose antennules, very reduced eyes—if at all present—, and biramous post-frontal appendages characterized by the presence of well-developed “fingers” on the inner side, called endites.

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: Nektobenthic, Epibenthic, Mobile
Feeding strategies: Predator, Suspension feeder
Ecological Interpretations:

Williams et al. (2007) considered bradoriids to be most likely epibenthic dwellers feeding on organic detritus or small non-biomineralized prey items; an ecology comparable to that of ostracods. These authors also pointed out that the large disparity of carapace shapes probably reflected a variety of different sub-niches and lifestyles. Their size and abundance would have made them in turn a common food source for larger animals, as indicated by coprolite composition (Vannier & Chen 2005).

References:

  • CARON, J. B. and JACKSON, D. A. 2008. Paleoecology of the Greater Phyllopod Bed community, Burgess Shale. Palaeogeography, Palaeoclimatology, Palaeoecology, 258, 222–256.
  • HOU, X., WILLIAMS, M., SIVETER, D. J., SIVETER, D. J., ALDRIDGE, R. J. and SANSOM, R. S. 2010. Soft-part anatomy of the Early Cambrian bivalved arthropods Kunyangella and Kunmingella : significance for the phylogenetic relationships of Bradoriida. Proceedings of the Royal Society B: Biological Sciences, 277, 1835–1841.
  • HOU, X., SIVETER, D. J., WILLIAMS, M., WALOSSEK, D., BERGSTRÖM, J., SMITH, M. P. and THOMAS, A. T. 1996. Appendages of 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. Palaeontology Newsletter, 32.
  • HUO, S. 1956. Brief notes on lower Cambrian Archaeostraca from Shensi and Yunnan. Acta Palaeontologica Sinica, 4, 1131–1145.
  • LI, Y. 1975. On the Cambrian ostracods with new material from Sichuan, Yunnan and Shaanxi, China. Professional Papers on Stratigraphy & Palaeontology, 2, 37–72.
  • MAAS, A., WALOSZEK, D. and MÜLLER, K. J. 2003. Morphology, ontogeny and phylogeny of the Phosphatocopina (Crustacea) from the Upper Cambrian ‘Orsten’ of Sweden. Fossil and Strata. Vol. 49. Taylor & Francis, Oslo.
  • MELNIKOVA, L. M. 1988. Nekotoryye bradoriidy (Crustacea) iz botomskogo yarusa vostochnogo Zabaykal’ya. Paleontologicheskiy Zhurnal, 1, 114–117.
  • MILLER, S. A. 1889. North American geology and palaeontology for the use of amateurs, students and scientists. Western Methodist Book Concern, Cincinnati.
  • QIAN, Y. and ZHANG, S. 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 CHEN, L. 1994. Cambrian palaeobiogeography of Bradoriida. Journal of Southeast Asian Earth Sciences, 9, 289–299.
  • SHU, D. G., VANNIER, J., LUO, H. L., CHEN, L., ZHANG, X. L. and HU, S. X. 1999. Anatomy and lifestyle of Kunmingella (Arthropoda, Bradoriida) from the Chengjiang fossil Lagerstätte (lower Cambrian; Southwest China). Lethaia, 32, 279–298.
  • SIVETER, D. J. and WILLIAMS, M. 1997. Cambrian bradoriid and phosphatocopid arthropods of North America. Special Papers in Palaeontology, 57, 67.
  • TOPPER, T. P., SKOVSTED, C. E., BROCK, G. A. and PATERSON, J. R. 2007. New bradoriids from the lower Cambrian Mernmerna Formation, South Australia: systematics, biostratigraphy and biogeography. Memoirs of the Association of Australasian Palaeontologists, 33, 67–100.
  • VANNIER, J. and CHEN, J. 2005. Early Cambrian food chain: New evidence from fossil aggregates in the Maotianshan Shale biota, SW China. PALAIOS, 20, 3–26.
  • WILLIAMS, M., SIVETER, D. J., POPOV, L. E. and VANNIER, J. M. C. 2007. Biogeography and affinities of the bradoriid arthropods: Cosmopolitan microbenthos of the Cambrian seas. Palaeogeography, Palaeoclimatology, Palaeoecology, 248, 202–232.
  • WRONA, R. 2009. Early Cambrian bradoriide and phosphatocopide arthropods from King George Island, West Antarctica: Biogeographic implications. Polish Polar Research, 30, 347–377.
  • ZHAI, D., WILLIAMS, M., SIVETER, D. J., HARVEY, T. H. P., SANSOM, R. S., GABBOTT, S. E., SIVETER, D. J., MA, X., ZHOU, R., LIU, Y. and HOU, X. 2019. Variation in appendages in early Cambrian bradoriids reveals a wide range of body plans in stem-euarthropods. Communications Biology, 2, 329.
  • ZHANG, W. 1974. Bradoriida. In Handbook of Stratigraphy and Palaeontology of Southwest China, Science Press, Beijing, 107–111 pp.
  • ZHANG, X.-G. 2007. Phosphatized Bradoriids (Arthropoda) from the Cambrian of China. Palaeontographica Abteilung A, 281, 93–173.
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