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The Cambrian Fossils of Chengjiang, China
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Table of Contents

Foreword ix Preface xi Part One Geological and Evolutionary Setting of the Biota 1 1 Geological time and the evolution of early life on Earth 3 2 The evolutionary significance of the Chengjiang biota 7 3 The discovery and study of the Chengjiang Lagerstatte 12 4 The distribution and geological setting of the Chengjiang Lagerstatte 20 5 The taphonomy and preservation of the Chengjiang fossils 26 6 The paleoecology of the Chengjiang biota 30 7 Cambrian Lagerstatten 35 Part Two Chengjiang Fossils 39 8 Algae 40 Fuxianospira gyrata Chen & Zhou, 1997 40 Megaspirellus houi Chen & Erdtmann, 1991 42 Sinocylindra yunnanensis Chen & Erdtmann, 1991 44 Yuknessia sp. of Chen & Erdtmann, 1991 46 9 Ctenophora 48 Galeactena hemispherica Ou et al., 2015 49 Maotianoascus octonarius Chen & Zhou, 1997 50 Batofasciculus ramificans Hou et al., 1999 52 Yunnanoascus haikouensis Hu et al., 2007 54 10 Porifera 56 Paradiagoniella xiaolantianensis Chen et al., 2014 56 Triticispongia diagonata Mehl & Reitner, 1993 58 Saetaspongia densa Mehl & Reitner, 1993 60 Leptomitus teretiusculus Chen et al., 1989 62 Choiaella radiata Rigby & Hou, 1995 64 Choia xiaolantianensis Hou et al., 1999 66 Allantospongia mica Rigby & Hou, 1995 68 Leptomitella conica Chen et al., 1989 70 Paraleptomitella dictyodroma Chen et al., 1989 72 Paraleptomitella globula Chen et al., 1989 74 Quadrolaminiella diagonalis Chen et al., 1990 76 11 Cnidaria 78 Archisaccophyllia kunmingensis Hou et al., 2005 78 Xianguangia sinica Chen & Erdtmann, 1991 80 12 Entoprocta 82 Cotyledion tylodes Luo & Hu, 1999 82 13 Phoronida 84 Iotuba chengjiangensis Chen & Zhou, 1997 84 14 Brachiopoda 86 Yuganotheca elegans Zhang et al., 2014 86 Heliomedusa orienta Sun & Hou, 1987 88 Longtancunella chengjiangensis Hou et al., 1999 90 Diandongia pista Rong, 1974 92 Lingulellotreta malongensis (Rong, 1974) 94 Lingulella chengjiangensis Jin et al., 1993 96 15 Annelida 98 Cambrosipunculus tentaculatus Huang et al., 2004 99 Maotianchaeta fuxianella Chen, 2004 100 16 Trochozoa of uncertain affinity 102 Ambrolinevitus maximus Jiang, 1982 103 Ambrolinevitus ventricosus Qian, 1978 104 Burithes yunnanensis Hou et al., 1999 106 Linevitus opimus Yu, 1974 108 Nectocaris pteryx Conway Morris, 1976 110 Wiwaxia papilio Zhang et al., 2015 112 17 Priapulida and relatives 114 Mafangscolex sinensis (Hou & Sun, 1988) 114 Maotianshania cylindrica Sun & Hou, 1987 116 Cricocosmia jinningensis Hou & Sun, 1988 118 Paraselkirkia sinica (Luo & Hou, 1999) 120 Corynetis brevis Luo & Hu, 1999 122 Sicyophorus rarus Luo & Hu, 1999 124 Palaeopriapulites parvus Hou et al., 1999 126 Eximipriapulus globocaudatus Ma et al., 2014 128 Xiaoheiqingella peculiaris Hu, 2002 130 Omnidens amplus Hou et al., 2006 132 Acosmia maotiania Chen & Zhou, 1997 134 Archotuba elongata (Luo & Hou, 1999) 136 18 Lobopodians 138 Paucipodia inermis Chen et al., 1995 138 Diania cactiformis Liu et al., 2011 140 Microdictyon sinicum Chen et al., 1989 142 Onychodictyon ferox Hou et al., 1991 144 Cardiodictyon catenulum Hou et al., 1991 146 Hallucigenia fortis Hou & Bergstroem, 1995 148 Luolishania longicruris Hou & Chen, 1989 150 Antennacanthopodia gracilis Ou & Shu, 2011 152 19 Anomalocaridids 154 Anomalocaris saron Hou et al., 1995 154 Amplectobelua symbrachiata Hou et al., 1995 156 Lyrarapax unguispinus Cong et al., 2014 158 Cucumericrus decoratus Hou et al., 1995 160 20 Euarthropoda 162 Isoxys paradoxus Hou, 1987 163 Isoxys auritus (Jiang, 1982) 164 Pectocaris spatiosa Hou, 1999 166 Shankouia zhenghei Chen et al., 2004 168 Chengjiangocaris longiformis Hou & Bergstroem, 1991 170 Fuxianhuia protensa Hou, 1987 172 Pseudoiulia cambriensis Hou & Bergstroem, 1998 174 Fortiforceps foliosa Hou & Bergstroem, 1997 176 Occacaris oviformis Hou, 1999 178 Forfexicaris valida Hou, 1999 180 Jianfengia multisegmentalis Hou, 1987 182 Tanglangia longicaudata Luo & Hu, 1999 184 Parapeytoia yunnanensis Hou et al., 1995 186 Haikoucaris ercaiensis Chen et al., 2004 188 Alalcomenaeus sp. of Tanaka et al., 2013 190 Leanchoilia illecebrosa (Hou, 1987) 192 Retifacies abnormalis Hou et al., 1989 194 Pygmaclypeatus daziensis Zhang et al., 2000 196 Squamacula clypeata Hou & Bergstroem, 1997 198 Urokodia aequalis Hou et al., 1989 200 Sinoburius lunaris Hou et al., 1991 202 Acanthomeridion serratum Hou et al., 1989 204 Cindarella eucalla Chen et al., 1996 206 Xandarella spectaculum Hou et al., 1991 208 Skioldia aldna Hou & Bergstroem, 1997 210 Saperion glumaceum Hou et al., 1991 212 Kuamaia lata Hou, 1987 214 Naraoia spinosa Zhang & Hou, 1985 216 Misszhouia longicaudata (Zhang & Hou, 1985) 218 Eoredlichia intermedia (Lu, 1940) 220 Kuanyangia sp. of Hou & Bergstroem, 1997 222 Yunnanocephalus yunnanensis (Mansuy, 1912) 224 Parapaleomerus sinensis Hou et al., 1999 226 Kwanyinaspis maotianshanensis Zhang & Shu, 2005 227 Kunmingella douvillei (Mansuy, 1912) 228 Kunyangella cheni Huo, 1965 230 Primicaris larvaformis Zhang et al., 2003 232 Branchiocaris? yunnanensis Hou, 1987 234 'Canadaspis' laevigata (Hou & Bergstroem, 1991) 236 Chuandianella ovata (Lee, 1975) 238 Clypecaris pteroidea Hou, 1999 240 Combinivalvula chengjiangensis Hou, 1987 242 Synophalos xynos Hou et al., 2009 244 Yunnanocaris megista Hou, 1999 246 21 Chaetognatha 248 Protosagitta spinosa Hu, 2002 248 22 Hemichordata 250 Galeaplumosus abilus Hou et al., 2011 251 23 Ambulacraria of uncertain affinity 252 Rotadiscus grandis Sun & Hou, 1987 253 Eldonia eumorpha (Sun & Hou, 1987) 254 Phlogites longus Luo & Hu, 1999 256 24 Chordata 258 Shankouclava anningense Chen et al., 2003 258 Myllokunmingia fengjiaoa Shu et al., 1999 260 Zhongxiniscus intermedius Luo & Hu, 2001 262 Cathaymyrus haikouensis Luo & Hu, 2001 263 25 Bilateria of uncertain affinity 264 Dinomischus venustus Chen et al., 1989 264 Facivermis yunnanicus Hou & Chen, 1989 266 Vetulocystis catenata Shu et al., 2004 268 Yunnanozoon lividum Hou et al., 1991 270 26 Vetulicolians 272 Heteromorphus confusus (Chen & Zhou, 1997) 272 Pomatrum ventralis Luo & Hu, 1999 274 Yuyuanozoon magnificissimi Chen et al., 2003 276 Beidazoon venustum Shu, 2005 278 Vetulicola cuneata Hou, 1987 280 27 Animals of uncertain affinity 282 Nidelric pugio Hou et al., 2014 282 Allonnia phrixothrix Bengtson & Hou, 2001 284 Stromatoveris psygmoglena Shu et al., 2006 286 28 Species recorded from the Chengjiang biota 288 29 Phylogenetic arrangement of chapters 292 References 293 Systematic Index 308 General Index 312

About the Author

Hou Xian-guang is former Director, Key Laboratory for Palaeobiology, Yunnan University, Kunming David J. Siveter is Professor Emeritus of Paleontology, University of Leicester Derek J. Siveter is Professor Emeritus of Earth Sciences, University of Oxford Richard J. Aldridge was Professor Emeritus and F.W. Bennett Professor of Geology, University of Leicester Cong Pei-yun is Professor of Paleobiology, Yunnan University, Kunming Sarah E. Gabbott is Professor of Paleobiology, University of Leicester Ma Xiao-ya is Professor of Paleobiology, Yunnan University, Kunming, and the Natural History Museum, London Mark A. Purnell is Professor of Paleobiology, University of Leicester Mark Williams is Professor of Paleobiology, University of Leicester

Reviews

The first edition of The Cambrian fossils of Chengjiang was a 'must have' book for palaeontologists. Now I am afraid you are all going to have to fork out for the second edition. After 13 years, this completely revised edition reflects the significant amount of research that has taken place over the last decade or more on the remarkable Chengjiang lagerstatte. The fossil material of the Chengjiang lagerstatte is of course stuff to drool over and the envy of palaeontologists who have to deal with the more common forms of hard-part preservation. Over 30 species have been added to the taxonomic list, reflecting a greater understanding of the diversity and taxonomy of this biota, which now boasts a total of over 250 species. A significant number of these are still 'floating around' waiting for their systematic identity card so the Chengjiang case is far from being closed. The discovery is now over 30 years old, being first found in 1984 by Hou Xian-Guang, who is still active and one of the authors of this new edition. Since the first edition was published, the site has been inscribed on the UNESCO World Heritage List (in 2012) as a 'globally outstanding example of a major stage in the history of life, representing a paleobiological window of great significance'. As such, the 515-520 Ma Chengjiang biota complements Canada's slightly younger, 505 Ma Burgess Shale biota. The latter is somewhat less diverse (c 120 known species) but has a similar ecological structure; consequently, the taxonomic similarities and differences between the two are of particular evolutionary interest and significance. This second edition of The Cambrian fossils of Chengjiang is much more than a mere update. It provides an overview of the lagerstatte and the rapidly expanding literature on all aspects of its geology, much of which is in Chinese and otherwise difficult to access. The format of the book is larger and many of the photos are even better than before. The book would be a fine present for any palaeontologist - even if you have to treat yourself! Reviewed by: Douglas Palmer "The work is authoritative and highly illustrated; the high-quality illustrations were, and are, an immensely important aspect of the work. They show just how beauti-fully preserved these soft-bodied animals are and how, with the requisite skills, this extraordinary detail can be illustrated. It is essential that this book be on every paleo-biologist's bookshelf." - Paul Seldon, Priscum Summer 2018 "Very much like its predecessor, this book is bound to become a standard reference thanks to its very well contextualized introduction and really complete overview of the Chengjiang biota. Whether you are a natural science teacher, a specialist of the Cambrian Explosion, have an interest in palaeontology and evolution of early life, or you just like the weird diversity of forms in Cambrian animals, then this book is for you. I would recommend it to all palaeontologists and libraries, this is a must-have!" - Vincent Perrier, Paleontology Association Newsletter, July 2018

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