Abstract
The origins and radiations of metazoans, bilaterians, deuterostomes and vertebrates have been recognized as the four most important milestone events in animal history and, conversely four of the toughest and puzzling questions in evolutionary biology. Among them, the first two occurred in the deep time of the Precambrian and have left little information in the fossil records. Fortunately, intensive investigation into the Early Cambrian Chengjiang faunas in the past two decades have revealed numerous significant data on the main episode of the Cambrian Explosion, particularly information on the various earliest-known deuterostomes, leading to a much better understanding of the last two events. The “first fish”Haikouichthys andMyllokunmingia have been proved the oldest-known vertebrates, and to be among the most primitive ones, representing a key transitional group from acraniates to craniates. Indeed, “Paleontological work of Shu and collaborators is revising our understanding of the early evolution of chordates”. Moreover, the discovery of the Phylum Vetulicolia may also throw new light on the origin of deuterostomes. This extinct group with simple gill slits might represent one of the roots in the deuterostome lineage, as the evidence of molecular biology and developmental biology has predicted. Here the first discovery of tiny vetulicolians from the Chengjiang Lagerstätte is reported. The evolutionary trend from the giant to the small might represent a special adaptation in early animal history. Since all vetulicolians share similar characters in their anterior section (pharynx), and on the basis of differences in their posterior section, a new classification system is proposed, with a new class, the Heteromorphida erected. As both the “root” group vetulicolians and the “top” group vertebrates in the early deuterostome lineage, as well as those members in between, such as vetulocystids, yunnanozoans, cephalochordates and urochordates, have been recovered, a relatively complete early deuterostome phylogeny, which is compatible with phylogenies of extant deuterostomes, is nearly mature.
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References
Shu, D., Guidance to reading “On the Origin of Species” by Ch. Darwin, in Chinese Translation of On the Origin of Species, Beijing: Peking University Press, 2005; Xian: Shaanxi People Press, 2001.
Finnerty, J. R., Martindale, M. Q., The evolution of Hox cluster: insights from outgroup, Curr. Opin. Genet. Dev., 1998, 8: 681–687.
Knoll, A., Carroll, S. B., Early animal evolution: emerging views from comparative biology and geology, Science, 1999, 284: 2129–2137.
Nielsen, C., Animal Evolution: interrelationships of living phyla, 2nd Ed., Oxford: Oxford University Press, 2001, 1–562.
Dawkins, R., The Ancestor’s Tale-A Pilgrimage to the Dawn of Life, New York: Weidenfeld & Nicolson, 2004, 1–528.
Ayala, F. J., Rzhetsky, A., Origin of metazoan phyla, Molecular clocks confirm palaeontological estimates. Proc. Natl. Acad. Sci, USA, 1998, 95: 606–611.
Signor, P. W., Lipps, J. H., Origin and Radiation of the Metazoa, in Origin and Early Evolution of the Metazoa (eds. Lipps, J. H., Signor, P. W.), New York: Plenum Press, 1992, 3–23.
Collins, A. G., Evaluating multiple alternative hypotheses for the origin of Bilateria, Proc. Natl. Acad. Sci. USA, 1998, 95: 15458–15463.
Halanych, K. M., Considerations for reconstructing metazoan history: signal, resolution, and hypothesis testing, Am. Zool., 1998, 38: 929–941.
Peterson, K. J., Butterfield, N. J., Origin of the Eumetazoa: Testing ecological prediction of molecular clock against the Proterozoic fossil record, PNAS, 2005, 102(27): 9547–9552.
Glaessner, M. F., The Dawn of Animal Life, Cambridge: Cambridge University Press, 1984.
Fedonkin, M., Vendian body fossils and trace fossils, in Early Life on Earth, Nobel Symposium No. 84, (ed. Bengtson, S.), New York: Columbia University, 1994, 370–388.
Conway Morris, S., The Ediacaran biota and early metazoan evolution, Geol. Mag., 1985, 122: 77–81.
Yuan, X., Xiao, S., Yin, L. et al. Doushantuo Fossils: Life on the Eve of Animal Radiation, Hefei: University of Science and Technology of China Press, 2002, 1–171.
Xiao, S., Zhang, Y., Knoll, A., Three-dimensional preservation of algae and animal embryos in a Neoproterozoic phosphorite, Nature, 1998, 391: 553–558.
Xiao, S., Yuan, X., Knoll, A., Eumetazoan fossils in terminal Proterozoic phosphorites. PNAS, 2000, 97(25): 13684–13689.
Chen, J., Bottjer, D., Oliveri, P. et al., Small Bilaterian Fossils from 40 to 55 Million Years before the Cambrian, Science, 2004, 305: 218–222.
Bengtson, S., Budd, G., Comment on “Small Bilaterian Fossils from 40 to 55 Million Years before the Cambrian”, Science, 2004, 305: 1291.
Garstang, W., The morphology of the Tunicata and its bearing on the phylogeny of the Chordata, J. of the Microscopical Society, 1928, 72: 51–87
Romer, A. S., The Vertebrate Body, Third ed., London: W.B. Saunders Company, 1964, 1–626.
Janvier, P., Early Vertebrates, Oxford: Clarendon, 1996.
Kardong, K., Vertebrates--Comparative Anatomy, Function, Evolution, Second ed., Boston: McGraw-Hill, 1998, 1–747.
Shu, D., Luo, H., ConwayMorris, S., et al., Early Cambrian vertebrates from South China, Nature, 1999, 402: 42–46.
Janvier, P., Catching the first fish, Nature, 1999, 402: 21–22
Shu, D., Chen, L., Mosaic evolution of the earliest-known vertebrates, Geosciences (in Chinese with English summery), 2000, 14: 315–322.
Shu, D., Conway Morris, S., Han, J., et al., Head and Backbone of the Cambrian vertebrateHaikouichthys, Nature, 2003, 421: 526–529.
Shu, D., A paleontological perspective of vertebrate origin, Chinese Science Bulletin, 2003, 48(8): 725–735.
Janvier, P., Vertebrate characters and the Cambrian vertebrates, Palevol, 2003, 2: 523–531.
Zhang, X. G., Hou, X., Evidence for a single median fin-fold and tail in the Lower Cambrian vertebrate,Haikouichthys ercaicunensis, Journal of Evolutionary Biology, 2004, 17: 1157–1161.
Halanych, K. M., The new view of animal phylogeny, Annual Reviews of Ecology and Evolutionary Systematics, 2004, 35: 229–256.
Schaeffer, B., Deuterostome monophyly and phylogeny, Evol. Biol., 1987, 21: 179–235.
Gee, H., Deuterostome phylogeny: the context for the origin and evolution of the vertebrates, in Major Events in Early Vertebrate Evolution: Palaeontology, Phylogeny, genetics and development (ed. Ahlberg, P. E.), London: Taylor & Francis, 2001, 1–14.
Shu, D., Conway Morris, S., Han, J., et al., Primitive deuterostomes from the Chengjiang Lagerstatte (Lower Cambrian, China), Nature 2001, 414: 419–424
Gee, H., On the vetulicolians, Nature, 2001, 414: 407–409
Conway Morris, S., Shu, D., Deuterostome Evolution, in McGraw-Hill yearbook of Science and Technology, New York: McGraw-Hill, 2003, 79–82.
Shu, D., Conway Morris, S., Han, J. et al., Ancestral echinoderms from the Chengjiang deposits of China, Nature 2004, 430: 422–428
Lacalli, T. C., Vetulicolians are they deuterostomes? chordates? BioEssays, 2002, 24: 208–211.
Chen, A., Feng, H., Zhu, M. et al., A new vetulicolian from the early Cambrian Chengjiang fauna in Yunnan of China, Acta Geologica. Sinica, 2003, 77(3): 281–287.
Valentine, J., On the Origin of Phyla, Chicago: Chicago University Press, 2004, 1–614.
Luo, H., Fu, X., Hu, S. et al., New vetulicolids from the Lower Cambrian Guanshan Fauna, Kunming, Acta Geologica Sinica, 2005. 79(1): 1–6.
Hou, X. G., Early Cambrian large bivalved arthropods from Chengjiang, eastern Yunnan, Acta Palaeontologica, 1987, 26(3): 286–298.
Hou, X. G., Ramskoeld, L., Bergstroem, J., Composition and preservation of the Chengjiang fauna—a Lower Cambrian soft-bodied biota, Zool. Scripta, 1991, 20: 395–411.
Chen, J., Zhou, G., Biology of the Chengjiang fauna, Bulletin of National Museum of Natural Science (Taichung), 1997, 10: 11–105.
Chen, J., The Dawn of Animal World, Nanjing: Jiangsu Science & Technology Publishing House, 2004, 1–366.
Chen, J., Dzik, J., Edgecombe, G. D. et al., A possible early Cambrian chordate, Nature, 1995, 377: 720–722.
Chen, J., Huang, D. Li, C. W., An Early Cambrian craniate-like chordate, Nature, 1999, 402: 518–521.
Dzik, J.,Yunnanozoon and ancestry of chordates, Acta Palaeont. Polonica, 1995, 40: 341–360.
Holland, H. D., Chen, J., Origin and early evolution of the vertebrates: new insights from advances in molecular biology, anatomy, and palaeontology, BioEssays, 2001, 23: 142–151.
Mallatt, J., Chen, J., Fossil sister group of craniates: Predicted and found, Journal of Morphology, 2003, 258(1): 1–31.
Shu, D., Zhang, X. L., Chen, L., Reinterpretation ofYunnanozoon as the earliest known hemichordate, Nature, 1996, 380: 428–430.
Smith, M. P., Sansom, I. J., Cochrane, D., The Cambrian origin of vertebrates, in Major Events in Early Vertebrate Evolution: Palaeontology, Phylogeny, genetics and development (ed. Ahlberg, P.), London: Taylor & Francis, 2001, 67–84.
Shu, D., Conway Morris, S., Zhang, Z. et al., A New Species of Yunnanozoan with Implications for Deuterostome Evolution, Science, 2003, 299: 1380–1384.
Shu, D., Conway Morris, S., Response to Comment on “A New Species of Yunnanozoan with Implications for Deuterostome Evolution”, Science, 2003, 300: 1372, 1372.
Steiner, M., Zhu, M., Zhao, Y. et al., Lower Cambrian Burgess shale-type fossil associations of south China, Palaeogeography, Palaeoclimatology, Palaeoecology, Special Issue, 2005, 220: 12–152.
Shu, D., Zhang, X. L., Chen, L., New advance in the Study of the Chengjiang fossil Lagerstatte, Progress in Geology of China (1993–1996), 30th IGC, Beijing, 1996, 42–46.
Shu, D., Conway Morris, S., Zhang, X. L. et al., A pipiscid-like fossil from the Lower Cambrian of South China, Nature, 1999, 400: 746–749.
Luo, H., Hu, S., Chen, L. et al., Early Cambrian Chengjiang fauna from Kunming region, China, Kunming: Yunnan Science and Technology Press, 1999, 1–129.
Winchell, C. J., Sullivan, J., Cameron, C. B. et al., Evaluating hypotheses of deuterostome phylogeny and chordate evolution with new LSU and SSU ribosomal DNA data, Molecular Biology and Evolution, 2002, 19(5): 762–776.
Holland, L. Z., Kene, M., William, N. A. et al., Sequence and embryonic expression of the amphioxusengrailed gene (AmphiEn): the metameric pattern of transcription resembles that of its segment-polarity homolog inDrosophila, Development, 1997, 124: 1723–1732.
Stollewerk, A., Schoppmeier, M., Damen, W. G., Involvement ofNotch andDelta genes in spider segmentation. Nature, 2003, 423: 863–865.
Jefferies, R. P. S., The ancestry of the vertebrates, London: Cambridge University Press, 1986.
Jefferies, R. P. S., A defense of the calcichordates, Lethaia, 1997. 30: 1–10.
Jefferies, R. P. S., The origin and early fossil history of the chordate acustio-lateralis system, with remarks on the reality of the echinoderm-hemichordate clade, in Major Events in Early Vertebrate Evolution: Palaeontology, Phylogeny, genetics and development (ed. Ahlberg, P.), London: Taylor & Francis, 2001, 40–66.
Ruppert, E. E., Cameron, C. B., Frick, J. F., Endostyle-like features of the dorsal epibranchial ridge of an enteropneust and the hypothesis of dorsal-ventral axis inversion in chordates, Invert. Biol., 1999, 118: 202–212.
Smith, A., Echinoderm roots, Nature 2004, 430: 411–412.
Bergstroem, J., Ideas on early animal evolution, in Early Life on Earth, Nobel Symposium No. 84 (ed. Bengtson, S.), New York: Columbia University, 1994, 460–466.
Briggs, D. G., Erwin, D. H., Collier, F. J., The Fossils of the Burgess Shale, Washington: Smithsonian Institution Press, 1994, 1–238.
Christen, R., Molecular phylogeny and the origin of Metazoa, in Early Life on Earth, Nobel Symposium No. 84 (ed. Bengtson, S.), New York: Columbia University, 1994, 467–474.
Conway Morris, S., Early metazoan evolution: First steps to an integration of molecular and morphological data, in Early Life on Earth, Nobel Symposium No. 84 (ed. Bengtson, S.), New York: Columbia University, 1994, 450–459.
Conway Morris, S., Cambrian Lagerstatten: their distribution and significance. Philosophical Transactions of the Royal Society of London B 1985, 311: 49–65.
Conway Morris, S., The Crucible of Creation: The Burgess Shale and the Rise of Animals, Cambridge: Cambridge Univ. Press, 1998, 1–242.
Valentine, J., The Cambrian explosion, in Early Life on Earth, Nobel Symposium No. 84 (ed. Bengtson, S.), New York: Columbia University, 1994, 401–411.
Chen, J., Huang, D., Peng, Q. et al. The first tunicate from the early Cambrian of South China, in Proceedings of the National Academy of Sciences of the USA, 2003, 100(14): 8314–8318.
Luo, H. L., Hu, S., Chen, L. Z., New Early Cambrian chordates from Haikou, Kunming, Acta Geologica Sinica, 2001, 75: 345–348.
Shu, D., Conway Morris, S., Zhang, X. L., A Pikaia-liks chordate from the Lower Cambrian of China, Nature, 1996, 384: 157–158.
Shu, D., Chen, L., Han, J. et al., The early Cambrian tunicate from South China, Nature, 2001, 411: 472–473.
Zhang, X. L., Shu, D., Li, Y. et al., New sites of Chengjiang fossils: crucial windows on the Cambrian explosion, J. Geol. Society, Lond., 2001, 158: 211–218.
Berrill, N. J., The Origin of Vertebrates, Oxford: Oxford University Press, 1955.
Bromham, L. D., Degnan, B. M., Hemichordate and deuterostome evolution: robust molecular phylogenetic support for a hemichordate + echinoderm clade, Evol. Dev., 1999, 1: 166–171.
Turner, S., Blieck, A. R. M., Nowlan, G. S., Cambrian-Ordovician vertebrates, in The Great Ordovician Biodiversity Event, IGCP410 (eds. Webby, B., Droser, M., Feist, R. et al.), New York: Columbia University Press, 2004, 327–335.
Briggs, D. G., Lieberman, B. S., Halgedah, S. L. et al., A new metazoan from the middle Cambrian of Utah and the nature of the Vetulicolia, Palaeontology, 2005, 48(4): 681–686.
Briggs, D. G., Fortey, R. A., Wonderful strife: Systematics, stem groups, and the phylogenetic signal of the Cambrian radiation. in Macroevolution: Diversity, Disparity, Contingency (eds. Vrba, E. S., Eldredge, N.), Kansas: The Paleontological Society, Lawrence, Supplement to Paleobiology, 2005, 31(2): 94–112.
Hou, X., Aldridge, R. J., Siveter, D. J. et al., New evidence on the anatomy and phylogeny of the earliest vertebrates. Proceedings of Royal Society of London, B, 2002, 269: 1865–1869.
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Shu, D. On the Phylum Vetulicolia. Chin.Sci.Bull. 50, 2342–2354 (2005). https://doi.org/10.1007/BF03183746
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DOI: https://doi.org/10.1007/BF03183746