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《地球科学与环境学报》[ISSN:1672-6561/CN:61-1423/P]

卷:

第31卷

期数:

2009年第02期

页码:

111-134

栏目:

基础地质与矿产地质

出版日期:

2009-06-15

文章信息/Info

Title:

Cambrian Explosion: Formation of Tree of Animals

文章编号:

1672-6561(2009)02-0111-24

作者:

舒德干¹; 2; 3

(1.西北大学 早期生命研究所,陕西 西安 710069; 2.西北大学 大陆动力学国家重点实验室, 陕西 西安 710069; 3.中国地质大学 地球科学与资源学院,北京 100083)

Author(s):

SHU De-gan¹; 2; 3

1.Early Life Institute, Northwest University, Xi'an 710069, Shaanxi, China; 2.State Key Laboratory of Continental Dynamics, Northwest University, Xi'an 710069, Shaanxi, China; 3.School of the Earth Sciences andResources, China University of Geosciences, Beijing 100083, China

关键词:

寒武纪大爆发; 动物树; 澄江化石库; 后口动物亚界; 古虫动物门; 脊椎动物起源

Keywords:

cambrian explosion;  tree of early animals;  Chengjiang fauna;  phylogeny of deuterostomes;  phylum vetulicolia;  origin of vertebrates

分类号:

Q91

DOI:

-

文献标志码:

A

摘要:

由“真动物”构成的动物界(不包括海绵)通常被划分为双胚层动物、原口动物和后口动物3个亚界。寒武纪大爆发经历了爆发的前奏—序幕—主幕3个阶段,其中后两个阶段处于早寒武世初期(依次以小壳化石的首次辐射和澄江动物群爆发为代表),另一个则发生在“寒武前夜”(即前寒武纪末期),以伊迪卡拉生物群为代表。这次独特的三幕式大爆发分步完成了动物形态演化谱系树(简写为TOA)的成型。已有化石证据显示,动物树的3大主体或3个亚界的起源及其早期辐射分别发生于寒武纪大爆发这3个主要阶段。澄江化石库中发现的早期后口动物亚界涵盖了该亚界中所有6大分支(棘皮类、半索类、头索类、尾索类、脊椎类和绝灭了的古虫类)的原始类群,澄江动物群时代标志着寒武纪大爆发的顶峰,完成了动物树框架的成型,从而宣告了寒武纪大爆发的基本终结。其学术重要性超越了中寒武世的布尔吉斯页岩,因为前者位于动物树形成的“源头”,而后者仅代表此后的“一段流程”。在动物演化历史上4个最具转折意义的重大创新事件(即多细胞动物起源、两侧对称动物起源、后口动物起源和脊椎动物起源)中,第一个发生在伊迪卡拉纪甚至更早的时期,其余3个则可通过梅树村化石群和澄江化石群观察到其完成的主体过程。已灭绝的古虫动物门,是初具鳃裂构造的原始分节动物,作为由原口动物向后口动物过渡的一个珍稀“缺环”,很可能代表着后口动物亚界的一个根。古囊类兼有两侧对称的古虫类和一些原始棘皮动物的镶嵌体征,最可能代表棘皮动物的一个根。在早寒武世动物演化谱系中,后口动物的“顶端类群”的代表是被誉为“第一鱼”的昆明鱼目,目前已知包括昆明鱼、海口鱼和钟健鱼,它们具对眼、脊索与串珠状脊椎软骨共存的原始脊椎,代表了真正的脊椎类或有头类始祖。相反,包括云南虫属和海口虫属的云南虫类,既没有成对的眼睛也没有可信的脊椎软骨,而且其背神经索也未扩大成脑,甚至不具备低等脊索动物的肌节,因此与真正的有头类或脊椎动物无关。其实,这类奇特动物与古虫类共享相似的躯体构形,应该代表着低等后口动物中的一个侧枝。自达尔文以来,进化生物学一直在动物世界的悠长演化历程中,努力追寻“人类远古由来”的一些重大形态学创新事件的证据,无疑,古虫动物门原始鳃裂构造的出现以及第一鱼之头颅和原始脊椎软骨的形成,应是这条历史长河中人们期待已久的两大里程碑。

Abstract:

The Kingdom Animalia is here divided into three subkingdoms: Diploblasta, Protostomia and Deuterostomia. Cambrian Explosion, including its first episode represented by the small skeletal fossils(SSFs)at the base of the Cambrian and the succeeding main episode represented by the Chengjiang faunas(CFs), together with its prelude represented by the Ediacaran faunas, have been recognized to give birth of the whole morphological tree of animals(or metazoans), for short the TOA. Its “seed” in Precambrian, some sort of protists from which the complete TOA have grown, remains unknown paleontologically. Ediacaran fauna was dominated by diploblasts(the “trunk” of the TOA)with a few possible stem-group triploblasts, the Early Cambrian in two phases explosively yielded almost all the major triploblastic crown-branches(Bilateria: the huge “crown” of the TOA), which include the other two Subkingdoms: first the extremely diverse protostomes in the Meishucunian Age and followed by a nearly entire lineage of early deuterostomes, i.e. the early or even the oldest representatives of all the six major groups in deuterostomes from the Chengjiang, including echinoderms, hemichordates, urochordates, cephalochordates and the vertebrates. As all the major branches of the Tree of Animals had been formed at the Chengjiang Age, the Cambrian Explosion reached its summit and also principally came to the end by the Early Cambrian. Among the four most significant milestone events of morphological origins and radiations in animal history, the first one(i.e. appearance of metazoans)took place in the Ediacaran Period or deeper times, and the other three can be practically seen from the Chengjiang and the Meishucunian windows. The newly discovered extinct Phylum Vetulicolia, which has primitively segmented body with simple gill slits in its anterior division, most probably represents one of the roots in deuterostome subkingdom. Sharing the mosaic basic features of both the bilateral vetulicolians and some primitive echinoderms homalozoans, the soft-bodied vetulocystids are best regarded as one of the roots of the extant pentamerous radial echinoderms. Standing on the “top” of the deuterostome super-branch in the early Cambrian TOA are the “the first fish” Myllokunmingia and Haikouichthys, which bear the definite paired eyes and the salient proto-vertebrae and then should represent the real root of the whole vertebrates or craniates. On the contrary, yunnanozoans, including Yunnanozoon and Haikouella, possess neither eyes and reliable vertebrae, nor myomeres, may have nothing to do with craniates or vertebrates. On the other hand, sharing a similar body-plan with vetulicolians, those enigmatic creatures might be best treated as a side-branch in the lower deuterostomes. The appearance of gill slits in Vetulicolians and the formation of the real brain and the most primitive vertebrae in “the First Fish” have been recognized as two of the most significant milestones in the evolution from protostomes to the vertebrates.

参考文献/References:

[1] Conway-Morris S. The Crucible of Creation: The Burgess Shale and the Rise of Animals[M].Oxford:Oxford University Press,1998.
[2] Cloud P E.Some Problems and Patterns of Evolution Exemplified by Fossil Invertebrates[J].Evolution,1948,2(4):322-350.
[3] Glaessner M F.New Fossils from the Base of the Cambrian in South Australia(Preliminary Account)[J].Transactions of the Royal Society of South Australia,1958,81:188-195.
[4] Glaessner M F.The Dawn of Animal Life[M].Cambridge:Cambridge University Press, 1984.
[5] Seilacher A.Vendozoa:Organismic Construction in the Proterozoic Biosphere[J].Lethaia,1989,22:229-239.
[6] McMenamin M A S.Vendian and Ediacaran[C]∥Selley R C,Cocks L R M,Plimer I R.Encyclopedia of Geology:Volume 4.Amsterdam:Elsevier,2005:371-381.
[7] McMenamin M A S,McMenamin D L S.The Emergence of Animal[M].New York:Columbia University Press,1990.
[8] Bengtson S,Conway-Morris S,Jell P A,et al.Early Cambrian Fossils from South Australia[J].Memoirs of the Association of Australasian Palaeontologists,1990,9:1-364.
[9] Bergström J.Precambrian Trace Fossils and the Rise of Bilateral Animals[J].Ichnos,1990,1:3-13.
[10] Fedonkin M A.Vendian Faunas and the Early Evolution of the Metazoa[C]∥Lipps J H,Signor P W.Origin and Early Evolution of the Metazoa.New York:Plenum Press,1992:87-129.
[11] Runnegar B.Evolution of the Earliest Animals[C]∥Schopf J W.Major Events in the History of Life.Boston: Jones and Bartlett,1992:65-93.
[12] Seilacher A.Vendobontia and Psammocorallia: Lost Con-
structions of Precambrian Evolution[J].Journal of the Geological Society,1992,149(4):607-613.
[13] Conway-Morris S.The Fossil Record and the Early Evolution of the Metazoa[J].Nature,1993,361:219-225.
[14] Fedonkin M A.Vendian Body Fossils and Trace Fossils[C]∥Bengtson S.Early Life on Earth: Nobel Symposium,Volume 84.New York: Columbia University Press,1994:370-388.
[15] Runnegar B.Proterozoic Eukaryotes: Evidence from Biology and Geology[C]∥Bengtson S.Early Life on Earth: Nobel Symposium,Volume 84.New York: Columbia University Press,1994:287-297.
[16] Grotzinger J P,Bowring S A,Saylor B Z,et al.Biostrati-graphic and Geochronologic Constraints on Early Animal Evolution[J].Science,1995,270:598-604.
[17] Collins A G.Evaluating Multiple Alternative Hypotheses for the Origin of Bilateria: an Analysis of 18S rRNA Molecular Evidence[J].Proceedings of the National Academy of Sciences of the United States of America,1998,95(26):15458-15463.
[18] Runnegar B.Understanding the Ediacaran Organisms: How to Proceed?[J].Abstracts with Programs-Geological Society of America,1998,30:147.
[19] Knoll A H,Carroll S B.Early Animal Evolution: Emerging Views from Comparative Biology and Geology[J].Science,1999,284:2129-2137.
[20] Erwin D H.The origin of Bodyplans[J].American Zoologists,1999,39:617-629.
[21] Conway-Morris S.The Cambrian “Explosion”:Slow-fuse or Egatonnage?[J].Proceedings of National Academy of Sciences of the United States of America,2000,97(9):4426-4429.
[22] Conway-Morris S.The Cambrian “Explosion” of Metazoans and Molecular Biology: Would Darwin be Satisfied?[J].International Journal of Developmental Biology,2003,47(7/8):505-515.
[23] Erwin D H,Davidson E H.The Last Common Bilaterian Ancestor[J].Development,2002,129: 3021-3032.
[24] Valentine J W.Prelude to the Cambrian Explosion[J].Annual Review of Earth and Planetary Sciences,2002,30:285-306.
[25] Valentine J W.On the Origin of Phyla[M].Chicago: Univer-sity of Chicago Press,2004.
[26] Waggoner B.The Ediacaran Biotas in Space and Time[J].Integrative Comparative Biology,2003,43:104-113.
[27] Benton M,Ayala F J.Dating the Tree of Life[J].Science,2003,300:1698-1700.
[28] Runnegar B.A Molecular-clock Date for the Origin of the Animal Phyla[J].Lethaia,1982,15:199-205.
[29] Runnegar B.The Cambrian Explosion: Animals or Fossils?[J].Journal of the Geological Society of Australia,1982,29:395-411.
[30] Signor P W,Lipps J H.Origin and Early Radiation of the Metazoa[C]∥Lipps J H,Signor P W.Origin and Early Evolution of the Metazoa.New York: Plenum Press,1992:3-23.
[31] Bengtson S.Protorozoic and Earliest Cambrian Skeleton Metazoans[C]∥Schopf W J,Klein C.The Protorozoic Biosphere-Amultidisciplinary Study.Cambridge: Cambridge University Press,1992:1017-1054.
[32] Qian Y,Bengtson S.Palaeontology and Biostratigraphy of the Early Cambrian Meishucun Stage in Yunnan Province,South China[J].Fossils and Strata,1989,24:1-156.
[33] Crimes T P.The Record of Trace Fossils Across the Proterozoic-Cambrian Boundary[C]∥Lipps J H.Signor P W.Origins and Early Evolution of Metazoa.New York:Plenum Press,1992:177-202.
[34] Fortey R A,Seilacher A.The Trace Fossil Cruziana Semiplicata and the Trilobite that Made It[J].Lethaia,1997,30:105-112.
[35] Conway-Morris S.The Burgess Shale Fauna and the Cambrian Explosion[J].Science,1989,246:339-346.
[36] Briggs D E G,Kear A J.Decay of the Lancelet Branchiostoma Lanceolatum(Cephalochordata): Implication for the Interpretation of Softtissue Preservation in Conodonts and Other Primitive Chordates[J].Lethaia,1994,26:275-287.
[37] Briggs D E G,Erwin D H,Collier F J.The Fossils of the Burgess Shale[M].Washington: Smithsonian Institution Press,1994.
[38] Shu D,Zhang X,Chen L.New Advance in the Study of the Chengjiang Fossil Lagerstatte[C]∥Geological Society of China.Progress in Geology of China(1993-1996): Papers to 30th IGC.Beijing: China Ocean Press,1996:42-46.
[39] Chen J,Zhou G.Biology of the Chengjiang Fauna[J].Bulletin of National Museum of Natural Science: Taichung,1997,10:11-105.
[40] 侯先光,杨·伯格斯琼,王海峰,等.澄江动物群:5.3亿年前的海洋动物[M].昆明:云南科技出版社,1999.
[41] Shu D.On the Phylum Vetulicolia[J].Chinese Science Bulletin,2005,50(20):2342-2354.
[42] 舒德干.澄江化石库中主要后口动物类群起源的初探[C]∥戎嘉余,方宗杰,周忠和,等.生物的起源、辐射与多样性演变——华夏化石记录的启示.北京:科学出版社,2006:109-124.
[43] 李国祥,Steiner M,钱 逸,等.华南寒武纪早期骨骼动物的爆发性辐射[C]∥戎嘉余,方宗杰,周忠和,等.生物的起源、辐射与多样性演变——华夏化石记录的启示.北京:科学出版社,2006:41-57.
[44] Grant S W F.Shell Structure and Distribution of Cloudina a Potential Index Fossil for the Terminal Proterozoic[J].American Journal of Science,1990,290A:261-294.
[45] Hua H,Pratt B R,Zhang L Y.Borings in Cloudina Shells: Complex Predator-prey Dynamics in the Terminal Neoproterozoic[J].Palaios,2003,18: 454-459.
[46] Grotzinger J P,Watters W A,Knoll A H.Calcified Metazoans in Thrombolite-stromatolite Reefs of the Terminal Proterozoic Nama Group,Namibia[J].Paleobiology,2000,26:334-339.
[47] 华 洪,陈 哲,张录易.后生动物骨骼化的起源[C]∥戎嘉余,方宗杰,周忠和,等.生物的起源、辐射与多样性演变——华夏化石记录的启示.北京:科学出版社,2006:29-39.
[48] Valentine J W.The Cambrian Explosion[C]∥Bengtson S.
Early Life on Earth: Nobel Symposium,Volume 84.New York: Columbia University Press,1994:401-411.
[49] Conway-Morris S.Evolution: Bringing Molecules into the Field[J].Cell,2000,100:1-11.
[50] Fortey R A,Jackson J,Strugnell J.Phylogenetic Fuses and Evolutionary Explosions; Conflicting Evidence and Critical Tests[C]∥Donoghue P C J,Smith M P.Telling the Evolutionary Time: Molecular Clocks and the Fossil Record.Boca Raton: CRC Press,2004:41-65.
[51] Sprigg R C.On the 1946 Discovery of the Precambrian Ediacara Fossil Fauna in South Australia[J].Earth Sciences History,1988,7:46-51.
[52] Fedonkin M A,Waggoner B M.The Late Precambrian Fossil Kimberella is a Mollusc-like Trilaterian Organism[J].Nature,1997,388:868-871.
[53] Xiao S,Zhang Y,Knoll A H.Three-dimensional Preservation of Algae and Animal Embryos in a Neoproterozoic Phosphorite[J].Nature,1998,391:553-558.
[54] Conway-Morris S.Ediacaran-like Fossils in Cambrian Bur-gess Shale-type Faunas of North America[J].Paleontology,1993,36:593-635.
[55] Shu D,Conway-Morris S,Han J,et al.Lower Cambrian Vendobionts from China and Early Diploblast Evolution[J].Science,2006,312:731-734.
[56] 舒德干,Conway-Morris S.澄江化石库中的双胚层动物新知[J].地学前缘,2006,13(6):251-227.
[57] Erwin D H.Molecular Clocks,Molecular Phylogenies and the Origin of Phyla[J].Lethaia,1989,22:251-257.
[58] Ayala F J,Rzhetsky A.Origin of Metazoan Phyla,Molecular Clocks Confirm Palaeontological Estimates[J].Proceedings of the National Academy of Sciences of the United States of America,1998,95:606-611.
[59] Donoghue P C J,Smith M P.Telling the Evolutionary Time: Molecular Clocks and the Fossil Record[M].Boca Raton: CRC Press,2004.
[60] Graur D,Martin W.Reading the Entrails of Chickens: Molecular Timescales of Evolution and the Illusion of Precision[J].Trends in Genetics,2004,20(2):80-86.
[61] Crimes T P.Trace Fossils[C]∥Cowie J W,Brasier M D.Oxford Monographs on Geology and Geophysics: Volume 12,The Precambrian-Cambrian Boundary.Oxford: Clarendon Press,1989:166-195.
[62] Rasmussen B,Bengtson S,Fletcher I R,et al.Discoidal Impressions and Trace-like Fossils More than 1 200 Million Years Old[J].Science,2002,296:1112-1115.
[63] McCall G J H.Earth Science Review:Volume 77,The Vendian(Ediacaran)in the Geological Record: Enigmas in Geology's Prelude to the Cambrian Explosion[M].Amsterdam:Elsevier,2006.
[64] Stanley S M.An Ecological Theory for the Sudden Origin of Multicellular Life in the Late Precambrian[J].Proceedings of the National Academy of Sciences of the United States of America,1973,70:1486-1489.
[65] McMenamin M A S.The Garden of Ediacara[J].Palaios,1986,1:178-182.
[66] Schopf J W.The Oldest Fossils and What They Mean[C]∥Schopf J W.Major Events in the History of Life.Boston:Jones and Bartlett,1992:29-63.
[67] Cloud P.Pre-Metazoan Evolution and the Origins of the Metazoa[C]∥Drake E T.Evolution and Environment.New Haven:Yale University Press,1968:1-72.
[68] Knoll A H.On the Age of the Doushantuo Formation[J].Acta Micropalaeontologica Sinica,1999,16:225-236.
[69] Canfield D E,Poulton S W,Narbonne G M.Late-neoproterozoic Deep-ocean Oxygenation and the Rise of Animal Life[J].Science,2007,315:92-95.
[70] Fike D A,Grotzinger J P,PrattL M,et al.Oxidation of the Ediacan Ocean[J].Nature,2006,444:744-747.
[71] Yin L,Zhu M,Knoll A,et al.Doushantuo Embryos Pre-served Inside Diapause Egg Cysts[J].Nature,2007,446:661-663.
[72] Valentine J W,Moores E M.Global Tectonics and the Fossil Record[J].Journal of Geology,1972,80:167-184.
[73] Knoll A H.Biological and Biogeochemical Preludes to the Ediacaran Radiation[C]∥Lipps J H,Signor P W.Origin and Early Evolution of the Metazoa.New York:Plenum Press,1992:53-84.
[74] Knoll A H,Walter M R.Latest Proterozoic Stratigraphy and Earth History[J].Nature,1992,356:673-678.
[75] Knoll A H.Neoproterozoic Evolution and Climate Change[C]∥Bengtson S.Early Life on Earth: Nobel Symposium,Volume 84.New York: Columbia University Press,1994:439-449.
[76] Maruyama S,Santosh M,Zhao D.Superplume,Superconti-nent,and Post-perovskite: Mantle Dynamics and Anti-plate Tectonics on the Core-mantle Boundary[J].Gondwana Research,2007,11:7-37.
[77] Grey K,Walter M R,Calver C R.Neoproterozoic Biotic Diversification: Snowball Earth or Aftermath of the Acraman Impact?[J].Geological Society of America,2003,31(5):459-462.
[78] Williamson D.Hybridization in the Evolution of Animal form and Lifecycle[J].Zoological Journal of the Linnean Society,2006,148:585-602.
[79] 达尔文.物种起源[M].舒德干,等译.北京:北京大学出版社,2005.
[80] Bowler P.Evolution: The History of an Idea[M].Berkeley: University of California Press,1989.
[81] Woese C R.Phylogenetic Trees,Whithermicrobiology?[J].Current Biology,1996,6:1060-1063.
[82] Müller W A.Developmental Biology[M].Heidelberg:Sprin- ger-Verlag,1995.
[83] Doolittle W F.Uprooting the Tree of Life[J].Scientific American,2000,282:90-95.
[84] Clegg M T,Gaut B S,Learn G H,et al.Rates and Patterns of Chloroplast DNA Evolution[J].Proceedings of the National Academy of Sciences of the United States of America,1994,91:6795-6801.
[85] Shu D.A Paleontological Perspective of Vertebrate Origin[J].Chinese Science Bulletin,2003,48(8):725-735.
[86] Shu D,Conway-Morris S,Han J,et al.Ancestral Echino-derms from the Chengjiang Deposits of China[J].Nature,2004,430:422-428.
[87] Brasier M D. Prokaryote Fossils[J].Encyclopedia of Geology,2005,4:363-370.
[88] Sun W.Early Multicellular Fossils[C]∥Bengston S.Early Life on Earth: Nobel Symposium,Volume 84.New York: Columbia University Press,1994:358-375.
[89] Xiao S.Precambrian Eukaryote Fossils[C]∥Selley R C,Cocks L R M,Plimer I R.Encyclopedia of Geology: Volume 4.Amsterdam: Elsevier,2005:354-363.
[90] Walter M R,Du R,Horodyski R.Coiled Carbonaceous-megafossils from the Middle Proterozoic of Jixian(Tianjin)and Montana[J].American Journal of Science,1990,290A: 133-148.
[91] 邢裕盛.中国震旦系[C]∥《国际地质学交流论文集》编写组.国际地质学交流论文集(二):地层学和古生物学.北京:地质出版社,1976:1-12.
[92] Sprigg R C.Early Cambrian(?)Jellyfish from the Flinders
Ranges,South Australia[J].Transactions of the Royal Society of South Australia,1947,71:212-223.
[93] Ford T D.Precambrian Fossils from the Charnwood Forest[J].Proceedings of the Yorkshire Geological Society,1958,31:211-217.
[94] Brasier M D,Hewitt R A,Brasier J.On the Late Precambrian-Early Cambrian Hartshill Formation of Warwickshire[J].Geological Magazine,1978,115:21-36.
[95] Richter R.Die Altesten Fossilien Sud-afrikas[J].Sencken-berg Lethaia,1955,36:243-289.
[96] Pflug H D.New Fossil Remains from the Nama Beds of South-west Africa[J].Paläontologische Zeitschrift,1966,40:14-25.
[97] Sokolov B S,Fedonkin M A,Ivanovskii A B.The Vendian
System:Regional Geology[M].Berlin: Springer,1990.
[98] Sistema V,Sokolov B S,Ivanovski A B.The Vendian System:
Paleontology[M].Berlin: Springer,1990.
[99] Hofmann H J.First Record of a Late Proterozoic Faunal Assemblage in the North American Cordillera[J].Lethaia,1981,14:303-310.
[100] Narbonne G M,Aitken J D.Ediacaran Fossils from the Sekwi Brook and Mackenzie Mountains,Yukon,Canada[J].Palaeontology,1990,33:945-980.
[101] Glaessner M F,Walter M R.New Precambrian Fossils from the Arumbera Sandstone,Northern Territory,Australia[J].Alcheringa,1975,1:59-69.
[102] Grey K,Griffin T J.King Leopold and Halls Creek Oro-gens: Local Sedimentary Succession Related to the Orogens[J].Memoir-Geological Survey of Western Australia,1990,3:249-255.
[103] Cruse T,Harris L B,Rasmussen B.Geological Note-The Discovery of Ediacaran Trace and Body Fossils in the Stirling Range Formation,Western Australia: Implications for Sedimentation and Deformation During the “Pan-African” Orogenic Cycle[J].Australian Journal of Earth Sciences,1993,40(3):293-296.
[104] Xiao S.Shen B,Zhou C,et al.A Uniquely Preserved Ediacaran Fossil with Direct Evidence for a Quilted Bodyplan[J].Proceedings of the National Academy of Sciences of the United States of America,2005,102:10227-10232.
[105] 袁训来,肖书海,周传明.新元古代陡山沱期真核生物的辐射[C]∥戎嘉余,方宗杰,周忠和,等.生物的起源、辐射与多样性演变——华夏化石记录的启示.北京:科学出版社,2006:13-27.
[106] 丁莲芳,李 勇,胡夏嵩,等.震旦纪庙河生物群[M].北京:地质出版社,1996.
[107] Grazhdankin D.Patterns of Distribution in Ediacaran Biotas and Facies Versus Biogeography and Evolution[J].Paleontology,2004,30(2):203-221.
[108] Wade M.Preservation of Soft-bodied Animals at Ediacara,South Australia[J].Lethaia, 1968,1:238-267.
[109] Wade M.Hydrozoa and Scyphozoa and Other Medusoids from the Precambrian Ediacara Fauna,South Australia[J].Palaeontology,1972,15:197-225.
[110] Jenkins R J F.Interpreting the Oldest Fossil Cnidarian[J].Palaeontographica Americana,1984,54:95-104.
[111] Gehling J G.A Cnidarian of Actinian-grade from the Ediacaran Pound Subgroup,South Australia[J].Alcheringa,1988,12:299-314.
[112] Jenkins R J F.Functional and Ecological Aspects of Ediacaran Assemblages[C]∥Lipps J H,Signor P W.Origins and Early Evolution of Metazoa.New York:Plenum Press,1992:131-176.
[113] Gehling J G,Narbonne G M,Anderson M M.The First Named Ediacaran Body Fossil,Aspidella Terranovica[J].Palaeontology,2000,43:427-456.
[114] Pflug H D.Zur Fauna der Nama-schichten in Sudwest-afrika,Bau und Systematische Zugchorogkeit[J].Palaeontologigraphica,1972,139:134-170.
[115] Seilacher A,Grazhdankin D,Legouta A.Ediacaran Biota: the Dawn of Animal Life in the Shadow of Giant Protists[J].Paleontological Research,2003,7(1):43-54.
[116] McMenamin M A S.The Garden of Ediacara: Discovering the First Complex Life[M].New York:Columbia University Press,1998.
[117] Narbonne G M.Modular Construction of Early Ediacaran
Complex Life Forms[J].Science,2004,305:1141-1144.
[118] Fedonkin M A.Promorphology of the Vendian Bilateria and the Problem of the Origin of Metamerism of Articulata[C]∥Sokolov B S.Problematiki Pozdnego Dokembriya I Paleozoya.Moscow:Nauka,1985:79-92.
[119] Gehling J G,Rigby J K.Long Expected Sponges from the Neoproterozoic Ediacara Fauna of South Australia[J].Journal of Palaeontology,1996,70:185-195.
[120] Li C W,Chen J,Hua T E.Precambrian Sponges with Cellular Structures[J].Science,1998,279:879-882.
[121] Zhang X L,Reitner J.A Fresh Look at Dickinsonia: Removing it from Vendobionta[J].Acta Geologica Sinica,2006,80:636-642.
[122] 陈均远.动物世界的黎明[M].南京:江苏科学技术出版社,2004.
[123] Xiao S,Yuan X,Knoll A H.Eumetazoan Fossils in Terminal Proterozoic Phosphorites[J].Proceedings of the National Academy of Sciences of the United States of America,2000,97:13684-13689.
[124] Yin L,Xiao S,Yuan X.New Observation on Spicule-like Structure from Doushantuo Phosphorites at Weng'an,Guizhou Province[J].Chinese Science Bulletin,2001,46:1828-1832.
[125] 袁训来,肖书海,尹磊明,等.陡山沱期生物群:早期动物辐射前夕的生命[M].合肥:中国科学技术大学出版社,2002.
[126] Chen J,Bottjer D,Gao F,et al.Small Bilaterian Fossils from 40 to 55 Million Years Before the Cambrian[J].Science,2004,305:218-222.
[127] Valentine J W.How Were Vendobiont Bodies Patterned[J].
Palaeobiology,2001,27:425-428.
[128] Budd G.A Cambrian Gilled Lobopod from Greenland[J].Nature,1993,364:709-711.
[129] Budd G.The Morphology of Opabinia Regalis and the Reconstruction of the Arthropod Stem-group[J].Lethaia,1996,29:1-14.
[130] Conway-Morris S,Peel J S.Articulated Halkieriids from the Lower Cambrian of North Greenland and Their Role in Early Protostome Evolution[J].Philosophical Transactions of the Royal Society B:Biological Sciences,1995,347:305-358.
[131] Williams M,Siveter D J,Peel J S.Isoxys(Arthropoda)from the Early Cambrian Sirius Passet Lagerstätte,North Greenland[J].Journal of Paleontology,1996,70:947-954.
[132] Budd G.The Morphology and Phylogenetic Significance of Kerygmachela Kierkegaardi Budd(Buen Formation,Lower Cambrian,N Greenland)[J].Transactions of the Royal Society of Edinburgh: Earth and Environmental Science,1999,89:249-290.
[133] Budd G,Jensen S.A Critical Reappraisal of the Fossil Record of the Bilaterian Phyla[J].Biological Review,2000,75:253-259.
[134] Bengtson S,Yue Z.Fossilized Metazoan Embryos from the Earliest Cambrian[J].Science,1997,277:1645-1648.
[135] Peel J S.Functional Morphology of the Class Helcionelloida nov,and the Early Evolution of the Mollusca[C]∥Simonetta A M,Conway-Morris S.The Early Evolution of Metazoa and the Significance of Problematic Taxa.Cambridge: Cambridge University Press,1991:157-177.
[136] Rozanov A Y,Zhuravlev A Y.The Lower Cambrian Fossil Record of the Soviet Union[C]∥Lipps J H,Signor P W.Origins and Early Evolution of Metazoa.New York:Plenum Press,1992:205-282.
[137] Bengtson S.The Cap-shaped Cambrian Fossil Maikhanella and the Relationship Between Coeloscleritophorans Andmolluscs[J].Lethaia,1992,25:401-420.
[138] Jiang Z.The Lower Cambrian Fossil Record of China[C]∥Lipps J H,Signor P W.Origins and Early Evolution of Metazoa.New York:Plenum Press,1992:311-333.
[139] Popov L Y.The Cambrian Radiation of Brachiopods[C]∥Lipps J H,Signor P W.Origins and Early Evolution of Metazoa.New York:Plenum Press,1992:399-423.
[140] Landing E.Lower Cambrian of Southeastern Newfound-land:Epeirogeny and Lazarus Faunas,Lithofacies-Biofacies Linkages,and the Myth of a Global Chronostratigraphy[C]∥Lipps J H,Signor P W.Origins and Early Evolution of Metazoa.New York:Plenum Press,1992:283-309.
[141] Feng W,Mu X,Sun W,et al.Microstructure of Early Cambrian Ramenta from China[J].Alcheringa,2002,26:9-17.
[142] Feng W,Sun W.Phosphate Replicated and Replaced Microstructure of Molluscan Shells from the Earliest Cambrian of China[J].Acta Paleontologica Polonica,2003,48:21-30.
[143] Li G,Xiao S.Micrina and Tannuolina(Tannuolidae)from the Lower Cambrian of Eastern Yunnan,South China[J].Journal of Paleontology,2004,78:900-913.
[144] 冯伟民.早寒武世单壳类软体动物最早的演化线系[C]∥戎嘉余,方宗杰,周忠和,等.生物的起源、辐射与多样性演变——华夏化石记录的启示.北京:科学出版社,2006: 59-72.
[145] Walcott C D.Cambrian Geology and Palaeontology II.3.Middle Cambrian Holothurians and Medusae[J].Smithsonian Miscellaneous Collection,1911,57:41-68.
[146] Walcott C D.Cambrian Geology and Paleontology II.5.Middle Cambrian Annelids[J].Smithsonian Miscellaneous Collections,1911,57:109-144.
[147] Whittington H B.Redescription of Marella Splendens(Trilobitoidea)from the Burgess Shale,Middle Cambrian,British Columbia[J].Bull Geol Surv Can,1971,209:1-24.
[148] Collins D H,Briggs D E G,Conway-Morris S.New Burgess Shale Fossils Sites Reveal Middle Cambrian Faunal Complex[J].Science,1983,222:163-167.
[149] Whittington H B.The Burgess Shale[M].New Haven:Yale University Press,1985.
[150] Gould S J.Wonderful Life: The Burgess Shale and the Nature of History[M].New York: W W Norton and Company,1989.
[151] Briggs D E G,Fortey R A.The Early Cambrian Radiation of Arthropods[C]∥ Lipps J H,Signor P W.Origins and Early Evolution of Metazoa.New York: Plenum Press,1992:335-373.
[152] Butterfield N J.A Reassessment of the Enigmatic Burgess Shale Fossil Wiwaxia Corrugate(Matthew)and Its Relationship to the Polychaete Canadia Spinosa Walcott[J].Paleobiology,1990,16:287-303.
[153] Butterfield N J.Leanchoilia Guts and the Interpretation of Three-dimensional Structures in Burgess Shale-type Fossils[J].Paleobiology,2002,28:155-171.
[154] Butterfield N J,Nicholas C J.Burgess Shale-type Preservation of Both Non-mineralizing and “Shelly” Cambrian Organisms from the Mackenzie Mountains,Northwestern Canada[J].Journal of Paleontology,1996,70:893-899.
[155] Conway-Morris S,Collins D H.Middle Cambrian Cteno-phores from the Stephen Formation,British Columbia,Canada[J].Philosophical Transactions of the Royal Society B:Biological Sciences,1996,35:279-308.
[156] Briggs D E G.The Arthropod Branchiocaris n.gen.,Middle Cambrian,Burgess Shale,British Columbia[J].Bulletin Geological Survey of Canada,1976,264:1-29.
[157] Conway-Morris S.Fossil Priapulid Worms[J].Special Papers in Palaeontology,1977,20:95.
[158] Conway-Morris S.A New Metazoan from the Cambrian Burgess Shale,British Columbia[J].Paleontology,1977,20:623-640.
[159] Conway-Morris S.Cambrian Lagerstatten: Their distribution and Significance[J].Philosophical Transactions of the Royal Society B:Biological Sciences,1985,311:49-65.
[160] Bergström J.Opabinia and Anomalocaris: Unique Cambrian “Arthropods”[J].Lethaia,1986,19:241-246.
[161] Collins D H.The “Evolution” of Anomalocaris and Its Classification in the Arthropod Class Dinocarida(nov.)and Order Radiodonta(nov.)[J].Journal of Paleontology,1996,70:280-293.
[162] Sprinkle J.Morphology and Evolution of Blastozoan Echinoderms[M].Cambridge:Harvard University Museum of Comparative Zoology Special Publication,1973.
[163] Conway-Morris S,Whittington H B.The Animals of Burgess Shale[J].Schientific American,1979,241:122-133.
[164] Paul C R C,Smith A.The Early Radiation and Phylogeny of Echinoderms[J].Biological Review,1984,59:443-481.
[165] 张文堂,侯先光.Naraoia在亚洲大陆的发现[J].古生物学报,1985,24(6):591-595.
[166] 侯先光.云南澄江早寒武世大型双瓣壳节肢动物[J].古生物学报,1987,26(3):286-298.
[167] 侯先光.云南澄江早寒武世三个新的大型节肢动物[J].古生物学报,1987,26(3):272-285.
[168] 侯先光.云南澄江早寒武世两个保存附肢的节肢动物[J].古生物学报,1987,26(3):236-256.
[169] 陈均远,侯先光,路浩之.早寒武世高足杯状稀珍海生动物——Dinomischus(Entoprocta)及其生态模式[J].古生物学报,1989,28(1):58-71.
[170] 陈均远,侯先光,路浩之.云南澄江下寒武统细丝海绵化石[J].古生物学报,1989,28(1):17-27.
[171] 侯先光,陈均远.云南澄江早寒武世节肢类与环节类中间性生物——Luolishania gen.nov.[J].古生物学报,1989,28(2):207-213.
[172] 侯先光,陈均远.云南澄江早寒武世带触手的蠕形动物——Facivermis gen.nov.[J].古生物学报,1989,28(1):32-41.
[173] 侯先光,陈均远,路浩之.云南澄江早寒武世节肢动物[J].古生物学报,1989,28(1):42-57.
[174] Shu D.Cambrian and Early Ordovician “Ostracoda”(Bradoriida)in China[J].Courier Forschung Institute Senckenberg,1990,123:315-330.
[175] Chen J,Erdtmann B D.Lower Cambrian Lagerstätte from
Chengjiang,Yunnan,China: Insights for Reconstructing Early Metazoan Life[C]∥Simonetta A M,Conway-Morris S.The Early Evolution of Metazoan and the Significance of Problematic Taxa.Cambridge:Cambridge University Press,1991:57-76.
[176] Jin Y,Wang H.Revision of the Lower Cambrian Brachiopod Heiliomedusa Sun and Hou,1987[J].Lethaia,1992,25(1):35-49.
[177] 舒德干,陈 苓,张兴亮,等.云南澄江化石库早寒武世KIN动物群[J].西北大学学报:自然科学版,1992,22(增刊):31-38.
[178] Hou X G,Bergström J.Palaeoscolecid Worms May be Nematomorphs Rather Than Annelids[J].Lethaia,1994,27:11-17.
[179] Shu D,Chen L.Cambrian Palaeobiologeography of Bradoriida[J].Journal of Southeast Asian Earth Sciences,1994,9:289-299.
[180] Chen J,Zhou G,Ramsköld L.The Cambrian Lobopodian Microdictyonsinicum[J].Bulletin of the National Museum of Natural Science,1995,5:1-93.
[181] Chen J,Zhou G,Ramsköld L.A New Early Cambrian Onychophoran Like Animal,Paucipodia gen.nov.,from the Chengjiang Fauna,China[J].Transactions of the Royal Society of Edinburgh: Earth Sciences,1995,85:275-282.
[182] Hou X G,Bergström J.Cambrian Lobopodians: Ancestors of Extant Onychophorans?[J].Zoological Journal of the Linnean Society,1995,114:3-19.
[183] Hou X G,Bergström J.Arthropods from the Lower Cambrian Chengjiang Fauna,Southwest China[J].Fossils and Strata,1997,45:1-116.
[184] Bergström J,Hou X.Chengjiang Arthropods and Their Bearing on Early Arthropod Evolution[C]∥Edgecombe G D.Arthropod Fossils and Phylogeny.New York:Columbia Univiversity Press,1998:151-184.
[185] Hou X G.New Rare Bivalved Arthropods from the Lower Cambrian Chengjiang Fauna,Yunnan,China[J].Journal of Paleontology,1999,73:102-116.
[186] Shu D,Geyer G,Chen L,et al.Redlichiacean Trilobites with Preserved Soft-parts from the Lower Cambrian Chengjiang Fauna,Beringaria[J].Special Issue,1995,2:203-241.
[187] Shu D,Zhang X,Geyer G.Anatomy and Systematic Affinities of Lower Cambrian Bivalved Arthropod Isoxys Auritus[J].Alcheringa,1995,19(4):333-342.
[188] Chen J,Dzik J,Edgecombe G D,et al.A Possible Early Cambrian Chordate[J].Nature,1995,377:720-722.
[189] Hou X G,Bergström J,Ahlberg P.Anomalocaris and Other Large Animals in the Lower Cambrian Chengjiang Fauna of Southwest China[J].Geologiska Foreningens i Stockholm Forhandligar,1995,117:163-183.
[190] Zhu M.Precambrian-Cambrian Trace Fossils from Eastern Yunnan: Implications for Cambrian Explosion[J].Bulletin of the National Museum of Natural Science,1997,10:275-312.
[191] 罗惠麟,胡世学,陈良忠,等.昆明地区早寒武世澄江动物群[M].昆明:云南科学技术出版社,1999.
[192] Zhang X L,Han J,Shu D.A New Arthropod Pygmaclypeatus Daziensis from the Early Cambrian Chengjiang Lagerstatte,South China[J].Journal of Paleontology,2000,74(5):979-982.
[193] Zhang X L,Shu D,Han J,et al.New Sites of Chengjiang Fossils: Crucial Windows on the Cambrian Explosion[J].Journal of the Geological Society,2001,158:211-218.
[194] Zhu M,Zhang J M,Li G.The Early Cambrian Chengjiang Biota: Quarries of Non-mineralized Fossils at Maotianshan and Ma'anshan,Chengjiang County,Yunnan Province,China[C]∥Peng S,Babcock L E,Zhu M.Cambrian System of China and Korea: Guide to field Excursions.Hefei:University of Science and Technology of China Press,2001:219-225.
[195] Zhang X L,Han J,Shu D.The First Occurrence of Burgess Shale Arthropod Sidneyia(S.sinica)in the Early Cambrian Chengjiang Lagerstatte and the Revision of Arthropod Urokodia[J].Alcheringa,2002,26:1-8.
[196] Zhang Z F,Han J,Zhang X L,et al.Pediculate Brachiopod Diandongia Pista from the Lower Cambrian of South China[J].Acta Geologica Sinica,2003,77:288-293.
[197] Zhang X L,Han J,Zhang Z F,et al.Reconsideration of the Supposed Naraoiid Larvae from the Early Cambrian Chengjiang Lagerstätte,South China[J].Palaeontology,2003,46:447-465.
[198] Zhang X G,Hou X,Emig C C.Evidence of Lophophores Fossils: Critical Window on the Cambrian Explosion[J].Journal of the Geological Society,2003,158:211-218.
[199] Han J,Zhang X L,Zhang Z F,et al.A New Platy-armored Wormfrom the Early Cambrian Chengjiang Lagerstätte,South China[J].Acta Geologica Sinica,2003,77(1):1-6.
[200] Zhang X L,Han J,Zhang Z F,et al.Redescription of the Chengjiang Arthropod Squamaculaclypeata Hou and Bergström,from the Lower Cambrian,South-west China[J].Palaeontology,2004,47(2):1-13.
[201] Zhang Z F,Han J,Zhang X L,et al.Soft Tissue Preservation in the Lower Cambrian Linguloid Brachiopod from South China[J].Acta Palaeontologica Polonica,2004,49:259-266.
[202] Zhang Z F,Shu D G,Han J,et al.Morpho-anatomical Differences of the Early Cambrian Chengjiang and Recent Linguloids and Their Implications[J].Acta Zoologica,2005,86(4):277-288.
[203] Han J,Shu D G,Zhang Z F,et al.The Earliest-known Ancestors of Recent Priapulomorpha from the Early Cambrian Chengjiang Lagerstätte[J].Chinese Science Bulletin,2004,49(17):1860-1868.
[204] Huang D,Vannier J,Chen J.Recent Priapulidae and Their Early Cambrian Ancestors: Comparisons and Evolutionary Significance[J].Geobios,2004,37:217-228.
[205] Liu J N,Shu D,Han J,et al.A Rare Lobopod with Well-preserved Eyes from Chengjiang Lagerstätte and Its Implications for Origin of Arthropods[J].Chinese Science Bulletin,2004,49(10):1063-1071.
[206] Han J,Zhang X L,Zhang Z F,et al.A New Theca-bearing Early Cambrian Worm from the Chengjiang Fossil Lagerstätte,China[J].Alcheringa,2006,30:1-10.
[207] Liu J N,Han J,Simonetta A M,et al.New Observations of the Lobopod-like Worm Facivermis from the Early Cambrian Chengjiang Lagerstatte[J].Chinese Science Bulletin,2006,51:363-385.
[208] Liu J N,Shu D G,Han J,et al.A Large Xenusiid Lobopod with Complex Appendages from the Lower Cambrian Chengjiang Lagerstatte[J].Acta Palaeontologica Polonica,2006,51(2):215-222.
[209] Zhang Z F,Shu D G,Han J,et al.New Data on the Rare Chengjiang(Lower Cambrian,South China)Linguloid Brachiopod Xianshanella Haikouensis[J].Journal of Paleontology,2006,80:203-211.
[210] Zhang Z F,Han J,Zhang X L,et al.Note on the Gut Preserved in the Lower Cambrian Lingulellotreta(Lingulata,Brachiopoda)from South China[J].Acta Zoologica(Stockholm),2007,88:65-70.
[211] Zhang Z F,Shu D G,Han J,et al.A Gregarious Lingulid Brachiopod Longtancunella Chengjiangensis from the Lower Cambrian,South China[J].Lethaia,2007,40:11-18.
[212] Zhang X G,Aldridge R J.Development and Diversification of Trunk Plates of the Lower Cambrian Lobopodians[J].Palaeontology,2007,50:401-415.
[213] Zhang Z F,Robson S,Emig C,et al.Early Cambrian Radiation of Brachiopods:a Perspective from South China[J].Gondwana Research,2008,14(1/2):241-254.
[214] Chen J,Huang D,Li C W.An Early Cambrian Craniate-like Chordate[J].Nature,1999,402:518-521.
[215] Shu D G,Zhang X,Chen L.Reinterpretation of Yunnanozoon as the Earliest Known Hemichordate[J].Nature,1996,380:428-430.
[216] Shu D G,Conway-Morris S,Zhang X L.A Pikaia-like Chordate from the Lower Cambrian of China[J].Nature,1996,384:156-157.
[217] Shu D G,Conway-Morris S,Zhang X,et al.A Pipiscid-like Fossil from the Lower Cambrian of South China[J].Nature,1999,400:746-749.
[218] Shu D G,Luo H,Conway-Morris S,et al.Early Cambrian Vertebrates from South China[J].Nature,1999,402:42-46.
[219] 舒德干,陈 苓.最早期脊椎动物的镶嵌演化[M].现代地质,2000,14(3):315-322.
[220] Shu D G,Chen L,Han J,et al.An Early Cambrian Tunicate from China[J].Nature,2001,411:472-473.
[221] Shu D G,Conway-Morris S,Han J,et al.Primitive Deuterostomes from the Chengjiang Lagerstatte(Lower Cambrian,China)[J].Nature,2001,414:419-424.
[222] Luo H,Hu S,Chen L.New Early Cambrian Chordates from Haikou,Kunming[J].Acta Geologica Sinica,2001,75(4):345-348.
[223] Holland H D,Chen J Y.Origin and Early Evolution of the Vertebrates: New Insights from Advances in Molecular Biology,Anatomy,and Palaeontology[J].BioEssays,2001,23:142-151.
[224] Hou X G,Aldridge R J,Siveter D J,et al.New Evidence on the Anatomy and Phylogeny of the Earliest Vertebrates[J].Proceedings of the Royal Society B: Biological Sciences,2002,269:1865-1869.
[225] Chen A,Feng H,Zhu M,et al.A New Vetulicolian from the Early Cambrian Chengjiang Fauna in Yunnan of China[J].Acta Geologica Sinica,

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收稿日期:2009-03-25
基金项目: 国家自然科学基金重点项目(40830208); 国家重点基础研究发展计划项目(2006CB806401); 教育部长江学者和创新团队发展计划项目(PSRIRT); 西北大学大陆动力学国家重点实验室项目
作者简介: 舒德干(1946-),男,湖北鄂州人,教授,博士研究生导师,从事古生物学和进化生物学研究。E-mail:elidgshu@nwu.edu.cn

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