A restudy of the Burgess Shale (Cambrian) arthropod Emeraldella brocki and reassessment of its affinities

Journal of Systematic Palaeontology

Volumn 10, Issue 2, 2012, Pages 361-383

  Stein, Martin ^a   Selden, Paul A ^a  

^a UNIVERSITY OF KANSAS   (United States)

Author keywords

'Arachnomorphs'; Artiopoda; Emeraldella brocki; Euarthropoda

Indexed keywords

ARTHROPOD; CAMBRIAN; CLADISTICS; FOSSIL RECORD; LIMB; MORPHOLOGY; PALEOBIOLOGY; PHYLOGENY; RECONSTRUCTION; TAXONOMY;

AGNOSTUS PISIFORMIS; ARTHROPODA; CRUSTACEA; PYGIDIUM; TRILOBITOMORPHA;

EID: 84858054991     PISSN: 14772019     EISSN: 14780941     Source Type: Journal    
DOI: 10.1080/14772019.2011.566634     Document Type: Article

References (82)

1. Bengtson, S. 2000. Teasing fossils out of shales with cameras and computers. Palaeontologia Electronica, 3, 1-14.
2. Bergström, J. & Hou, X. 2005. Early Palaeozoic nonlamellipedian arthropods. Pp. 73-93 in S. Koenemann & R. A. Jenner (eds) Crustacean Issues 16. Crustacea and Arthropod Relationships. Taylor & Francis, Boca Raton.
3. Bergström, J., Hou, X., Zhang, X. & Clausen, S. 2008. A new view of the Cambrian arthropod Fuxianhuia. GFF, 130, 189-201.
4. Beurlen, K. 1934. Die Pygaspiden, eine neue Crustacean-(Entomostracen)- gruppe aus den Mesosaurier führenden Iraty-Schichten Brasiliens. Paläontologische Zeitschrift, 16, 122-138.
5. Bremer, K. 1994. Branch support and tree stability. Cladistics, 10, 295-304.
6. 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, 1-24.
7. Briggs, D. E. G., Bruton, D. L. & Whittington, H. B. 1979. Appendages of the arthropod Aglaspis spinifer (Upper Cambrian, Wisconsin) and their significance. Palaeontology, 22, 167-180.
8. Bruton, D. L. & Nakrem, H. A. 2005. Enrollment in a Middle Ordovician agnostoid trilobite. Acta Palaeontologica Polonica, 50, 441-448. (Pubitemid 41410654)
9. Bruton, D. L. & Whittington, H. B. 1983. Emeraldella and Leanchoilia, two arthropods from the Burgess Shale, Middle Cambrian, British Columbia. Philosophical Transactions of the Royal Society, Series B, 300, 553-582.
10. Budd, G. E. 2008. Head structure in upper stem-group euarthropods. Palaeontology, 51, 561-573. (Pubitemid 351699596)
11. Chen, J., Edgecombe, G. D., Ramsköld, L. & Zhou, G. 1995. Head segmentation in Early Cambrian Fuxianhuia: implications for arthropod evolution. Science, 268, 1339-1343.
12. Chen, J., Zhou, G., Zhu, M. & Yeh, K. 1996. The Chengjiang Biota. A unique window of the Cambrian Explosion. The NationalMuseum of Natural Sciences, Taichung, Taiwan, 222 pp.
13. Chen, J., Edgecombe, G. D. & Ramsköld, L. 1997. Morphological and ecological disparity in naraoiids (Arthropoda) from the Early Cambrian Chengjiang Fauna, China. Records of the Australian Museum, 49, 1-24.
14. Chen, J., Waloszek, D. & Maas, A. 2004. A new 'greatappendage' arthropod from the Lower Cambrian of China and homology of chelicerate chelicerae and raptorial anteroventral appendages. Lethaia, 37, 3-20. (Pubitemid 38688606)
15. Cisne, J. L. 1981. Triarthrus eatoni (Trilobita): anatomy of its exoskeletal, skeletomuscular, and digestive systems. Palaeontographica Americana, 9, 99-142.
16. 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. (Pubitemid 39364616)
17. Cotton, T. J. & Fortey, R. A. 2005. Comparative morphology and relationships of the Agnostida. Crustacean Issues 16. Crustacea and Arthropod Relationships, Pp. 95-113 in S. Koenemann & R. A. Jenner (eds) Crustacean Issues 16. Crustacea and Arthropod Relationships. Taylor & Francis, Boca Raton.
18. Dzik, J. & Lendzion, K. 1988. The oldest arthropods of the East European Platform. Lethaia, 21, 29-38.
19. Edgecombe, G. D. & Ramsköld, L. 1999. Relationships of Cambrian Arachnata and the systematic position of Trilobita. Journal of Paleontology, 73, 263-287. (Pubitemid 29178241)
20. Farris, J. S., Albert, V.A., Källersjö, M., Lipscomb, D. & Kluge, A. G. 1996. Parsimony jackknifing outperforms neighbourjoining. Cladistics, 12, 99-181.
21. Fortey, R. A. 1990. Ontogeny, hypostome attachment and trilobite classification. Palaeontology, 33, 529-576.
22. Fortey, R. A. 2001. Trilobite systematics: the last 75 years. Journal of Paleontology, 75, 1141-1151. (Pubitemid 34267768)
23. Fortey, R. A. & Theron, J. N. 1994. A new Ordovician arthropod, Soomaspis, and the agnostid problem. Palaeontology, 37, 841-861.
24. Goloboff, P. A., Farris, J. S. & Nixon, K. 2008. TNT, a free program for phylogenetic analysis. Cladistics, 24, 774-786.
25. Hall, J. 1838. Descriptions of two species of trilobites belonging to the genus Paradoxides. American Journal of Science, 13, 139-142.
26. Haug, C., Haug, J. T., Waloszek, D., Maas, A., Frattigiani, R. & Liebau S. 2009a. New methods to document fossils from lithographic limestones of southern Germany and Lebanon. Palaeontologia Electronica, 12, 1-12.
27. Haug, J. T., Maas, A. & Waloszek, D. 2009b. † Henningsmoenicaris scutula, †Sandtorpia vestrogothiensis gen. et sp. nov. and heterochronic events in early crustacean evolution. Earth and Environmental Science Transactions of the Royal Society of Edinburgh, 100, 311-350.
28. Hendricks, J. R. & Lieberman, B. S. 2008. New phylogenetic insights into the Cambrian radiation of arachnomorph arthropods. Journal of Paleontology, 82, 585-594. (Pubitemid 351730277)
29. Hesselbo, S. P. 1992. Aglaspidida (Arthropoda) from the Upper Cambrian of Wisconsin. Journal of Paleontology, 66, 885-923. (Pubitemid 23389276)
30. Hicks, H. 1875. On the succession of the ancient rocks in the vicinity of St. David's, Pembrokeshire, with special reference to those of the Arenig and Llandeilo groups, and their fossil contents. Quarterly Journal of the Geological Society of London, 31, 167-195.
31. Hou, X. 1987a. Three newlarge arthropods from Lower Cambrian Chengjiang, easternYunnan. Acta Palaeontologica Sinica, 26, 272-285.
32. Hou, X. 1987b. Two new arthropods from Lower Cambrian, Chengjiang, eastern Yunnan. Acta Palaeontologica Sinica, 26, 236-256.
33. Hou, X. & Bergström, J. 1991. The arthropods of the Lower Cambrian Chengjiang fauna, with relationships and evolutionary significance. Pp. 179-187 in A. M. Simonetta & S. Conway Morris (eds) The early evolution of Metazoa and the significance of problematic taxa. Cambridge University Press, Cambridge.
34. Hou, X. & Bergström, J. 1997. Arthropods of the Lower Cambrian Chengjiang Fauna, southwest China. Fossils and Strata, 45, 1-116. (Pubitemid 127654986)
35. Hou, X., Chen, J. & Lu, H. 1989. Early Cambrian newarthropods from Chengjiang, Yunnan. Acta Palaeontologica Sinica, 28, 42-57.
36. Hou, X., Ramsköld, L. & Bergström, J. 1991. Composition and preservation of the Chengjiang fauna - a Lower Cambrian soft-bodied biota. Zoologica Scripta, 20, 395-411.
37. Hou, X., Aldridge, R. J., Bergström, J., Siveter, D. J., Siveter, D. J. & Fen, X. 2004. The Cambrian Fossils of Chengjiang, China. The Flowering of Early Animal Life. Blackwell, Oxford, 233 pp.
38. Hou, X., Clarkson, E.N. K., Yang, J., Zhang, X., Wu, G. & Yuan, Z. 2009. Appendages of early Cambrian Eoredlichia (Trilobita) from the Chengjiang biota, Yunnan, China. Earth and Environmental Science Transactions of the Royal Society of Edinburgh, 99, 213-223.
39. Hughes, N. C., Haug, J. T. & Waloszek, D. 2008. Basal euarthropod development: a fossil based perspective. Pp. 281-298 in A. Minelli & G. Fusco (eds) Evolving pathways: key themes in evolutionary developmental biology. Cambridge University Press, Cambridge.
40. Jell, P. A. 2003. Phylogeny of early Cambrian trilobites. Special Papers in Palaeontology, 70, 45-57.
41. Lagebro, L., Stein, M. & Peel, J. S. 2009. A new ?lamellipedian arthropod from the early Cambrian Sirius Passet Fauna of North Greenland. Journal of Paleontology, 83, 820-825.
42. Lin, J. 2006. Taphonomy of naraoiids (Arthropoda) from the Middle Cambrian Kaili Biota, Guizhou Province, South China. Palaios, 21, 15-25. (Pubitemid 43257403)
43. Liu, Y., Hou, X. & Bergström, J. 2007. Chengjiang arthropod Leanchoilia illecebrosa (Hou, 1987) reconsidered. GFF, 129, 263-272. (Pubitemid 350042037)
44. Lu, Y. 1940. On the ontogeny and phylogeny of Redlichia intermedia Lu (sp. nov.). Bulletin of the Geological Society of China, 20, 333-342.
45. Maas, A., Waloszek, D., Chen, J., Braun, A., Wang, X. & Huang, D. 2004. Phylogeny and life habits of early arthropods - Predation in the Early Cambrian sea. Progress in Natural Science, 14, 158-166.
46. Maddison, W. P. & Maddison, D. R. 2010. Mesquite: a modular system for evolutionary anaylsis. Version 2.73 [updated at http://mesquiteproject.org], accessed 10 September 2010.
47. Manton, S. M. 1977. The Arthropoda. Habits, functional morphology, and evolution. Oxford University Press, Oxford, 528 pp.
48. Müller, K. J. 1983. Crustacea with preserved soft parts from the Upper Cambrian of Sweden. Lethaia, 16, 93-109.
49. Müller, K. J. & Walossek, D. 1987. Morphology, ontogeny, and life habit of Agnostus pisiformis from the Upper Cambrian of Sweden. Lethaia, 19, 1-124.
50. Novozhilov, N. I. 1960. Klass Merostomoidea Størmer, 1944, Chlenistonogie, Trilobitoobraznye i Rakoobraznye [Arthropoda, Trilobitomorpha and Crustacea]. Osnovy Paleontologii, 195-197.
51. Paterson, J. R., Edgecombe, G. D., García-Bellido, D. C., Jago, J. B. & Gehling, J. G. 2010. Nektaspid arthropods from the lower Cambrian Emu Bay Shale lagerstätte, South Australia, with a reassessment of lamellipedian relationships. Palaeontology, 53, 377-402.
52. Raasch, G. O. 1939. Cambrian Merostomata. Geological Society of America Special Paper, 16, 1-146.
53. Ramsköld, L. & Edgecombe, G. D. 1996. Trilobite appendage structure - Eoredlichia reconsidered. Alcheringa, 20, 269-276.
54. Ramsköld, L., Chen, J., Edgecombe, G. D. & Zhou, G. 1996. Preservational folds simulating tergite junctions in tegopeltid and naraoiid arthropods. Lethaia, 29, 15-20.
55. Ramsköld, L., Chen, J., Edgecombe, G. D. & Zhou, G. 1997. Cindarella and the arachnate clade Xandarellida (Arthropoda, Early Cambrian) from China. Transactions of the Royal Society of Edinburgh: Earth Sciences, 88, 19-38.
56. Raymond, P. E. 1920. The appendages, anatomy and relationships of trilobites. Memoirs of the Connecticut Academy of Arts and Sciences, 7, 1-169.
57. Rominger, C. 1887. Description of primordial fossils from Mt. Stephens, N. W. Territory of Canada. Proceedings of the Academy of Natural Sciences, Philadelphia, 1887, 12-19.
58. Scholtz, G. & Edgecombe, G. D. 2005. Heads, Hox and the phylogenetic position of trilobites. Pp. 139-165 in S. Koenemann & R. A. Jenner (eds) Crustacean Issues 16. Crustacea and Arthropod Relationships. Taylor & Francis, Boca Raton.
59. Simonetta, A.M. 1964. Osservazioni sugli artropodi non trilobiti della 'Burgess Shale' (Cambriano medio). III. Contributo. Monitore zoológico italiano, 72, 215-231.
60. Stein, M. 2010. A new arthropod from the Early Cambrian of North Greenland with a 'great appendage' like antennula. Zoological Journal of the Linnean Society, 158, 477-500.
61. Stein, M., Waloszek, D. & Maas, A. 2005. Oelandocaris oelandica and the stem lineage of Crustacea. Pp. 55-72 in S. Koenemann & R. A. Jenner (eds) Crustacean Issues 16. Crustacea and Arthropod Relationships. Taylor & Francis, Boca Raton.
62. Stein, M., Waloszek, D., Maas, A., Haug, J. T. & Müller, K. J. 2008. The stem crustacean Oelandocaris oelandica Müller re-visited. Acta Palaeontologica Polonica, 53, 461-484.
63. Størmer, L. 1944. On the relationships and phylogeny of fossil and recent Arachnomorpha. Skrifter Utgitt av Det Norske Videnskaps-Akademi i Oslo. I. Matematisk-Naturvetenskapelig Klasse, 1944 (5), 1-158.
64. Størmer, L. 1959. Trilobitoidea. Pp. O23-O37 in R. C. Moore (ed.) Treatise on Invertebrate Paleontology, Part. O, Arthropoda 1. Geological Society of America and University of Kansas Press, Lawrence.
65. Wahlenberg, G. 1821. Petrificata Telluris Svecanae. Nova acta Regiae Societatis scientiarum upsaliensis, 8, 1-116.
66. Walch, J. E. I. 1771. Die Naturgeschichte der Versteinerungen zur Erläuterung der Knorrischen Sammlung von Merkwürdigkeiten der Natur. Nürnberg.
67. Walcott, C. D. 1911. Cambrian geology and paleontology II. Middle Cambrian Merostomata. Smithsonian Miscellaneous Collections, 57, 17-40.
68. Walcott, C. D. 1912. Cambrian geology and paleontology II. Middle Cambrian Branchiopoda, Malacostraca, Trilobita and Merostomata. Smithsonian Miscellaneous Collections, 57, 145-229.
69. Walcott, C. D. 1918. Geological explorations in the Canadian Rockies. Explorations and fieldwork of the Smithsonian Institution in 1917. Smithsonian Miscellaneous Collections, 68, 4-20.
70. Walossek, D. & Müller, K. J. 1990. Upper Cambrian stemlineage crustaceans and their bearing upon the monophyletic origin of Crustacea and the position of Agnostus. Lethaia, 23, 409-427.
71. Waloszek, D., Chen, J., Maas, A. & Wang, X. 2005. Early Cambrian arthropods-new insights into arthropod head and structural evolution. Arthropod Structure & Development, 34, 189-205. (Pubitemid 40725688)
72. Waloszek, D., Maas, A., Chen, J. & Stein, M. 2007. Evolution of cephalic feeding structures and the phylogeny of Arthropoda. Palaeogeography, Palaeoclimatology, Palaeoecology, 254, 273-287. (Pubitemid 47432685)
73. Whittington, H. B. 1974. YohoiaWalcott and Plenocaris n. gen., arthropods from the Burgess Shale, Middle Cambrian, British Columbia. Geological Survey of Canada Bulletin, 231, 1-27.
74. Whittington, H. B. 1975. Trilobites with appendages from the Middle Cambrian, Burgess Shale, British Columbia. Lethaia, 4, 97-136.
75. Whittington, H. B. 1977. TheMiddleCambrian trilobite Naraoia, Burgess Shale, British Columbia. Philosophical Transactions of the Royal Society of London. Series B, 280, 409-443.
76. Whittington, H. B. 1980. Exoskeleton, moult stage, appendage morphology, and habits of the Middle Cambrian trilobite Olenoides serratus. Palaeontology, 23, 171-204.
77. Whittington, H. B. 1981. Rare arthropods from the Burgess Shale, Middle Cambrian, British Columbia. Philosophical Transactions of the Royal Society, Series B, 292, 329-357.
78. Whittington, H. B. & Almond, J. E. 1987. Appendages and habits of the Upper Ordovician trilobite Triarthrus eatoni. Philosophical Transactions of the Royal Society of London, Series B, 317, 1-46.
79. Zhang, W. & Hou, X. 1985. Preliminary notes on the occurrence of the unusual trilobite Naraoia in Asia. Acta Palaeontologica Sinica, 24, 591-595.
80. Zhang, X. & Shu, D. 2005. A new arthropod from the Chengjiang Lagerstätte, Early Cambrian, southern China. Alcheringa, 29, 185-194. (Pubitemid 41622700)
81. Zhang, X., Han, J. & Shu, D. 2000. A new arthropod Pygmaclypeatus daziensis from the Early Cambrian Chengjiang Lagerstätte, South China. Journal of Paleontology, 74, 979-983.
82. Zhang, X., Shu, D. & Erwin, D. H. 2007. Cambrian naraoiids (Arthropoda): morphology, ontogeny, systematics, and evolutionary relationships. Journal of Paleontology, 81, 1-52. (Pubitemid 47529079)