Phylogeny and evolutionary significance of vermiform animals from the Early Cambrian Chengjiang Lagerstätte
Tóm tắt
The Early Cambrian Chengjiang Biota of China has preserved many species of vermiform animals in exquisite detail, which provide a unique window into the origin and early evolution of different vermiform phyla. This article reviews and discusses the forty-five Chengjiang vermiform animals discovered so far, and specifically comments on the phylogeny, modes of life, evolutionary significance as well as existing debates concerning the Chengjiang priapulid and lobopodian phyla. This article analyzes existing problems with Chengjiang vermiform morphological research and highlights various factors that affect the final preserved fossil morphology, including taphonomy and shape change caused by vermiform movement, as well as proposing that Anningvermis multispinosa and Corynetis brevis are synonymous. Chengjiang vermiform animals displayed a great diversity of morphologies and ecological roles, which is significant for understanding the origin and evolution of early metazoans; the potential for future research is enormous but needs to be broadened by interdisciplinary work involving taphonomy, geochemistry, cladistics, and genetics.
Tài liệu tham khảo
Collins D. Misadventures in the Burgess Shale. Nature, 2009, 460: 952–953
Zhang W T, Hou X G. Preliminary notes on the occurrence of the unusual trilobite Naraoia in Asia (in Chinese with English abstract). Acta Palaeontol Sin, 1985, 24: 591–595
Sun W G, Hou X G. Early Cambrian worms from Chengjiang, Yunnan, China: Maotianshania gen. nov (in Chinese with English abstract). Acta Palaeontol Sin, 1987, 26: 299–305
Hou X G, Bergström J, Wang H F, et al. The Chengjiang Fauna: Exceptionally Well-preserved Animals from 530 Million Years Ago (in Chinese). Kunming: Yunnan Science and Technology Press, 1999. 170
Hou X G, Aldridge R J, Bergström J, et al. The Cambrian Fossils of Chengjiang, China: The Flowering of Early Animal Life. Oxford: Blackwell Publishing, 2004. 233
Hou X G, Sun W G. Discovery of Chengjiang fauna at Meishucun, Jinning, Yunnan (in Chinese with English abstract). Acta Palaeontol Sin, 1988, 27: 1–12
Hou X. G, Bergström J. Palaeoscolecid worms may be nematomorphs rather than annelids. Lethaia, 1994, 27: 11–17
Luo H L, Hu S X, Chen L Z, et al. Early Cambrian Chengjiang Fauna from Kunming Region, China (in Chinese). Kunming: Yunnan Science and Technology Press, 1999. 129
Han J, Zhang Z F, Shu D G. Discovery of the proboscis on Tylotites petiolaris (in Chinese). Northwestern Geol, 2003, 36: 87–93
Huang D Y, Vannier J, Chen J Y. Anatomy and lifestyles of Early Cambrian priapulid worms exemplified by Corynetis and Anningvermis from the Maotianshan Shale (SW China). Lethaia, 2004, 37: 21–33
Chen J Y. The Dawn of Animal World (in Chinese). Nanjing: Jiangsu Science and Technology Press, 2004. 366
Chen L Z, Luo H L, Hu S X, et al. Early Cambrian Chengjiang Fauna in East Yunnan, China (in Chinese). Kunming: Yunnan Science and Technology Press, 2002. 199
Huang D Y, Vannier J, Chen J Y. Recent Priapulidae and their Early Cambrian ancestors: Comparisons and evolutionary significance. Geobios, 2004, 37: 217–228
Han J, Zhang X L, Zhang Z F, et al. A new platy-armored worm from the Early Cambrian Chengjiang Lagerstätte, South China. Acta Geol Sin, 2003, 77: 1–6
Han J, Shu D G, Zhang, Z F, et al. The earliest-known ancestors of Recent Priapulomorpha from the Early Cambrian Chengjiang Lagerstätte. Chin Sci Bull, 2004, 49: 1860–1868
Hou X G, Berström J, Yang J. Distinguishing anomalocaridids from arthropods and priapulids. Geol J, 2006, 41: 259–269
Han J, Liu J N, Zhang Z F, et al. Trunk ornament on the palaeoscolecid worms Cricocosmia and Tabelliscolex from the Early Cambrian Chengjiang deposits of China. Acta Palaeontol Pol, 2007, 52: 423–431
Huang D Y. Early Cambrian Worms from SW China: Morphology, Systematics, Lifestyles and Evolutionary Significance. Doctor Dissertation. Lyon: Université Claude Bernard Lyon 1, 2005
Ma X Y, Hou X G, Siveter D J, et al. A new priapulid-like worm from the Early Cambrian Chengjiang Lagerstätte, China. In: Abstracts of the 51st Annual Meeting of the Palaeontological Association, 2007 Dec 16–19, Uppsala. Palaeontological Association Newsletter, 2007, 66: 43
Maas A, Huang D Y, Chen J Y, et al. Maotianshan-Shale nemathelminths: Morphology, biology, and the phylogeny of Nemathelminthes. Palaeogeogr Palaeoclimatol Palaeoecol, 2007, 254: 288–306
Whittard W F. Palaeoscolex piscatorum gen. et sp. nov., a worm from the Tremadocian of Shropshire. Q J Geol Soc Lond, 1953, 109: 125–136
Conway Morris S. The cuticular structure of a 495-Myr-old type species of the fossil worm Palaeoscolex, P. piscatorum (?Priapulida). Zool J Linn Soc, 1997, 119: 69–82
Chen J Y, Zhou G Q. Biology of the Chengjiang Fauna. Bull Natl Mus Nat Sci Taiwan, 1997, 10: 11–106
Harvey T H P, Dong X P, Donoghue C J. Are palaeoscolecids ancestral ecdysozoans? Evol Dev, 12: 177–200
Huang D Y, Chen J Y, Vannier J. Discussion on the systematic position of the Early Cambrian priapulomorph worms. Chin Sci Bull, 2006, 51: 243–249
Han J, Hu S X. Response to “Discussion on the systematic position of the Early Cambrian priapulomorph worms”. Chin Sci Bull, 2006, 51: 250–256
Dong X P, Donoghue P C J, Cunningham J A, et al. The anatomy, affinity, and phylogenetic significance of Markuelia. Evol Dev, 2005, 7: 468–482
Hou X G, Chen J Y. Early Cambrian Arthropod-Annelid intermediate sea animals, Luolishania gen. nov. from Chengjiang, Yunnan (in Chinese with English abstract). Acta Palaeontol Sin, 1989, 28: 207–213
Ma X Y, Hou X G, Bergström J. Morphology of Luolishania longicruris (Lower Cambrian, Chengjiang Lagerstätte, SW China) and the phylogenetic relationships within lobopodians. Arthropod Struct Dev, 2009, 38: 271–291
Liu J N, Shu D G, Han J, et al. A rare lobopod with well-preserved eyes from Chengjiang Lagerstätte and its implications for origin of arthropods. Chin Sci Bull, 2004, 49: 1063–1071
Chen J Y, Hou X G, Lu H Z. Early Cambrian netted scale-bearing worm-like sea animal (in Chinese with English abstract). Acta Palaeontol Sin, 1989, 28: 1–16
Hou X G, Ramsköld L, Bergström J. Composition and preservation of the Chengjiang fauna—A Lower Cambrian soft-bodied biota. Zool Scr, 1991, 20: 395–411
Liu J N, Shu D G, Han J, et al. The lobopod Onychodictyon from the Lower Cambrian Lagerstätte revisited. Acta Palaeontol Pol, 2008, 53: 285–292
Hou X G, Bergström J. Cambrian lobopodians-ancestors of extant onychophorans? Zool J Linn Soc, 1995, 114: 3–19
Chen J Y, Zhou G Q, Ramsköld L. A new Early Cambrian Onychophoran-like animal, Paucipodia gen. nov., from the Chengjiang fauna, China. Trans R Soc Edinb-Earth Sci, 1995, 85: 275–282
Hou X G, Ma X Y, Zhao J, et al. The lobopodian Paucipodia inermis from the Lower Cambrian Chengjiang fauna, Yunnan, China. Lethaia, 2004, 37: 235–244
Liu J N, Shu D G, Han J, et al. Morpho-anatomy of the lobopod Megadictyon cf. haikouensis from the Early Cambrian Chengjiang Lagerstätte, South China. Acta Zoologica, 2007, 88: 279–288
Liu J N, Shu D G, Han J, et al. A large Xenusiid lobopod with complex appendages from the Lower Cambrian Chengjiang Lagerstätte. Acta Palaeontol Pol, 2006, 51: 215–222
Dzik J, Krumbiegel G. The oldest’ onychophoran’ Xenusion: A link connecting phyla? Lethaia, 1989, 22: 169–181
Bergström J, Hou X G. Cambrian Onychophora or Xenusians. Zool Anz, 2001, 240: 237–245
Budd G. Tardigrades as ’stem-Group Arthropods’: The Evidence from the Cambrian Fauna. Zool Anz, 2001, 240: 265–279
Huang D Y, Chen J Y, Vannier J, et al. Early Cambrian sipunculan worms from southwest China. Proc R Soc B-Biol Sci, 2004, 271: 1671–1676
Chen J Y, Huang D Y. A possible Lower Cambrian Cheatognath (arrow worm). Science, 2002, 298: 187
Hu S X. Taphonomy and palaeoecology of the Early Cambrian Chengjiang Biota from Eastern Yunnan, China. Berliner Paläobiologische Abhandlungen, 2005, 7: 1–197
Hou X G, Cong P Y, Li Y Z. On the taphonomy and phylogenetic relationships of yunnanozoans (in Chinese with English abstract). Acta Palaeotol Sin, 2009, 48: 402–413
Chapman G. The Hydrostatic Skeleton in the Invertebrates. Biol Rev, 1958, 33: 338–371
Dornbos S Q, Chen J Y. Community palaeoecology of the early Cambrian Maotianshan Shale biota: Ecological dominance of priapulids worms. Palaeogeogr Palaeoclimataol Palaeoecol, 2008, 258: 200–212
Zhang X G, Hou X G, Bergström J. Early Cambrian priapulid worms buried with their lined burrows. Geol Mag, 2006, 143: 743–748
Hammond R A. The burrowing of Priapulus caudatus. J Zool Lond, 1970, 162: 469–480
Ruppert E E, Barnes R D. Invertebrate Zoology, 6th ed. New York: Sanders College Publishing, 1996. 1100
Han J, Zhang Z F, Liu J N. Evidence of Priapulid Scavenging from the Early Cambrian Chengjiang Deposits, Southern China. Palaios, 2007, 22: 691–694