24


Journal of Paleontology 92(1):14–25


assigned to total-group Scalidophora to indicate their uncertain positions within Scalidophora; Form B might represent at least a new species with closer affinities to E. rarus. All putative scalidophoran animals were recovered from


the Cambrian Fortunian Stage of South China only very recently. Since their diversity in the small shelly faunas is highly underestimated, the current research highlights the significance of Orsten-type Lagerstätten in probing the morphology and tax- onomy of the oldest known scalidophoran animals. Accordingly, it provides key information on the origin and early evolution of Scalidophora and Cycloneuralia. To summarize, in uncovering the origin and early evolution of Cycloneuralia, the Orsten-type Lagerstätten are progressively equal in systematic and evolutionary importance to Burgess Shale–type Lagerstätten.


Acknowledgments


This work was supported by the Chinese Ministry of Science and Technology 973 Project (2013CB837100), the National Natural Science Foundation of China (41572007), and the Youth Innovation Promotion Association, Chinese Academy of Sciences (2016283). Two anonymous reviewers provided constructive suggestions for this paper.


References


Aguinaldo, A.M.A., Turbeville, J.M., Linford, L.S., Rivera, M.C., Garey, J.R., Raff, R.A., and Lake, J.A., 1997, Evidence for a clade of nematodes, arthropods and other moulting animals: Nature, v. 387, p. 489–493.


Ahlrichs, W.H., 1995, Ultrastruktur und Phylogenie von Seison nebaliae (Grube 1859) und Seison annulatus (Claus 1876): Hypothesen zu phylogenetischen Verwandschaftsverhältnissen innerhalb der Bilateria: Göttingen, Cuvillier, 310 p.


Brusca, R.C., and Brusca, G.J., 2003, Invertebrates: Sunderland, Massachusetts, Sinauer Associates, 936 p.


Budd, G.E., 2001, Why are arthropods segmented?: Evolution & Development, v. 3, p. 332–342.


Butterfield, N.J., Balthasar, U., and Wilson, L.A., 2007, Fossil diagenesis in the Burgess Shale: Palaeontology, v. 50, p. 537–543.


Butterfield, N.J., 2003, Exceptional fossil preservation and the Cambrian Explosion: Integrative and Comparative Biology, v. 43, p. 166–177.


Chen, J., 2004, The Dawn of Animal World: Nanjing, Jiangsu Science and Technology Press, 366 p.


Cheng, G., Peng, F., Duan, B., and Dong, X.-P., 2011, Internal structure of Cambrian fossil embryo Markuelia revealed in the light of synchrotron radiation x-ray tomographic microscopy: Acta Geologica Sinica, v. 85, p. 81–90.


Conway Morris, S., 1989, Burgess Shale faunas and the Cambrian Explosion: Science, v. 246, p. 339–346.


Conway Morris, S., and Chen, M., 1991, Cambroclaves and paracarinachitids, early skeletal problematica from the lower Cambrian of South China: Palaeontology, v. 34, p. 357–397.


Dong, X.-P., Donoghue, P.C.J., Cheng, H., and Liu, J.B., 2004, Fossil embryos from the middle and late Cambrian Period of Hunan, South China: Nature, v. 427, p. 237–240.


Donoghue, P.C.J., and Purnell, M.A., 2009, Distinguishing heat from light in debate over controversial fossils: BioEssays, v. 31, p. 178–189.


Duan, B., Dong, X.-P., and Donoghue, P.C.J., 2012, New palaeoscolecid worms from the Furongian (upper Cambrian) of Hunan, South China: Is Markuelia an embryonic palaeoscolecid?: Palaeontology, v. 55, p. 613–622.


Gaines, R.R., Kennedy, M., and Droser, M., 2005,Anew hypothesis for organic preservation of Burgess Shale taxa in the middle Cambrian Wheeler Formation, House Range, Utah: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 220, p. 193–205.


Gaines, R.R., Briggs, D.E.G., and Zhao, Y.L., 2008, Cambrian Burgess Shale-type deposits share a common mode of fossilization: Geology, v. 36, p. 755–758.


Harvey, T.H.P., Dong, X.-P., and Donoghue, P.C.J., 2010, Are palaeoscolecids ancestral ecdysozoans?: Evolution & Development, v. 12, p. 177–200.


Haug, J.T., Maas, A., Waloszek, D., Donoghue, P.C.J., and Bengtson, S., 2009, A new species of Markuelia from the middle Cambrian of Australia,


in Paterson, J.R., and Laurie, J.R., eds., Cambro-Ordovician Studies III: Memoirs of the Association of Australasian Palaeontologists: Canberra, Association of Australasian Palaeontologists, v. 37, p. 303–313.


He, T.G., and Xie, Y.S., 1989, Some problematic small shelly fossils from the Meishucunian of the lower Cambrian in the western Yangtze Region: Acta Micropalaeontologica Sinica, v. 6, p. 111–127.


He, Y.X., and Yang, X.H., 1986, Early Cambrian coelenterates from Nanjiang, Sichuan: Bulletin of the Chengdu Institute of Geology and Mineral Resources, v. 7, p. 31–43.


Li, Z., 1984, The discovery and its significance of small shelly fossils in Hexi area, Xixiang, Shaanxi: Geology of Shaanxi, v. 2, p. 73–77.


Lemburg, C., 1995, Ultrastructure of the introvert and associated structures of the larvae of Halicryptus spinulosus (Priapulida): Zoomorphology, v. 115, p. 11–29.


Liu, Y., Xiao, S., Shao, T., Broce, J., and Zhang, H., 2014, The oldest known priapulid-like scalidophoran animal and its implications for the early evolution of cycloneuralians and ecdysozoans: Evolution & Development, v. 16, p. 155–165.


Maas, A., Braun, A., Dong, X.-P., Donoghue, P.C.J., Müller, K.J., Olempska, E., Repetski, J.E., Siveter, D.J., Stein, M., and Waloszek, D., 2006, The ‘Orsten’—More than a Cambrian Konservat-Lagerstätte yielding exceptional preservation: Palaeoworld, v. 15, p. 266–282.


Maas, A., Waloszek, D., Haug, J.T., and Müller, K.J., 2007a, A possible larval roundworm from the Cambrian ‘Orsten’ and its bearing on the phylogeny of Cycloneuralia: Memoirs of the Association of Australasian Palaeonto- logists, v. 34, p. 499–519.


Maas, A., Huang, D., Chen, J., Waloszek, D., and Braun, A., 2007b, Maotianshan-Shale nemathelminths: Morphology, biology, and the phylogeny of Nemathelminthes: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 254, p. 288–306.


Merriman, J.A., 1981, Cuticular structures of the priapulid Halicryptus spinulosus: A scanning electron microscopical study: Zoomorphology, v. 97, p. 285–295.


Missarzhevsky, V.V., 1973, Konodontoobraznye organizmy iz pogranichnykh sloev kembriya i dokembriya Sibirskoj platformy i Kazakhstana [Conodont- shaped organisms from Precambrian–Cambrian boundary strata of the Siberian Platform and Kazakhstan]: Trudy Instituta Geologii i Geofiziki SO AN SSSR, v. 49, p. 53–57.


Missarzhevsky, V.V., 1974, Novye dannye o drevnejshikh okamenelostyakh rannego kembriya Sibirskoj platformy [New data on the oldest fossils of the early Cambrian of the Siberian Platform], in Zhuravleva, I.T., and Rozanov, A.Y., eds., Biostratigrafiya i paleontologiya nizhnego kembriya Evropy i severnoj zii [Biostratigraphy and Palaeontology of the Lower Cambrian of Europe and Northern Asia]: Moscow, Nauka, p. 179–189.


Müller, K.J., 1985, Exceptional preservation in calcareous nodules: Philoso- phical Transactions of the Royal Society of London, Series B, Biological Sciences, v. 311, p. 67–73.


Müller, K.J., and Hinz-Schallreuter, I., 1993, Palaeoscolecid worms from the middle Cambrian of Australia: Palaeontology, v. 36, p. 549–592.


Müller, K.J., and Walossek, D., 1985, A remarkable arthropod fauna from the upper Cambrian ‘Orsten’ of Sweden: Transactions of the Royal Society of Edinburgh, v. 76, p. 161–172.


Müller, K.J., and Walossek, D., 1991, ‘Orsten’ arthropods–small in size but of great impact on biological and phylogenetic interpretations: Geologiska Föreningen i Stockholm Förhandlingar, v. 113, p. 88–90.


Neuhaus, B., 2013, Kinorhyncha (= Echinodera), in Schmidt-Rhaesa, A., ed., Handbook of Zoology: Gastrotricha, Cycloneuralia and Gnathifera, Volume 1: Nematomorpha, Priapulida, Kinorhyncha, Loricifera: Berlin/Boston, Walter de Gruyter GmbH, p. 181–348.


Neuhaus, B., and Higgins, R.P., 2002, Ultrastructure, biology, and phylogenetic relationships of Kinorhyncha: Integrative and Comparative Biology, v. 42, p. 619–632.


Neuhaus,B., Kristensen,R.M., andLemburg, C., 1996, Ultrastructure of the cuticle of the Nemathelminthes and electron microscopical localization of chitin: Verhandlungen der Deutschen Zoologischen Gesellschaft, v. 89, p. 221.


Neuhaus, B., Kristensen, R.M., and Peters, W., 1997, Ultrastructure of the cuticle of Loricifera and demonstration of chitin using gold-labelled wheat germ agglutinin: Acta Zoologica (Stockholm), v. 35, p. 193–213.


Nielsen, C., 2012, Animal Evolution: Interrelationships of the Living Phyla: Oxford, Oxford University Press, 402 p.


Peng, S., Babcock, L.E., and Cooper, R.A., 2012, The Cambrian Period, in Gradstein, F.M., Ogg, J.G., Schmitz, M., and Ogg, G., eds., Geological Time Scale 2012: Oxford, Elsevier, p. 437–488.


Qian, Y., 1977, Hyolitha and some problematica from the lower Cambrian Meishucun Stage in central and southwestern China: Acta Palaeontologica Sinica, v. 16, p. 255–275.


Missarzhevsky, V.V., 1983, Stratigrafiya drevnejshikh tolshch fanerozoya Anabarskogo massiva [Stratigraphy of the oldest Phanerozoic strata of the Anabar Massif]: Sovetskaya Geologiya, v. 1983, p. 62–73.


Page 1  |  Page 2  |  Page 3  |  Page 4  |  Page 5  |  Page 6  |  Page 7  |  Page 8  |  Page 9  |  Page 10  |  Page 11  |  Page 12  |  Page 13  |  Page 14  |  Page 15  |  Page 16  |  Page 17  |  Page 18  |  Page 19  |  Page 20  |  Page 21  |  Page 22  |  Page 23  |  Page 24  |  Page 25  |  Page 26  |  Page 27  |  Page 28  |  Page 29  |  Page 30  |  Page 31  |  Page 32  |  Page 33  |  Page 34  |  Page 35  |  Page 36  |  Page 37  |  Page 38  |  Page 39  |  Page 40  |  Page 41  |  Page 42  |  Page 43  |  Page 44  |  Page 45  |  Page 46  |  Page 47  |  Page 48  |  Page 49  |  Page 50  |  Page 51  |  Page 52  |  Page 53  |  Page 54  |  Page 55  |  Page 56  |  Page 57  |  Page 58  |  Page 59  |  Page 60  |  Page 61  |  Page 62  |  Page 63  |  Page 64  |  Page 65  |  Page 66  |  Page 67  |  Page 68  |  Page 69  |  Page 70  |  Page 71  |  Page 72  |  Page 73  |  Page 74  |  Page 75  |  Page 76  |  Page 77  |  Page 78  |  Page 79  |  Page 80  |  Page 81  |  Page 82  |  Page 83  |  Page 84  |  Page 85  |  Page 86  |  Page 87  |  Page 88  |  Page 89  |  Page 90  |  Page 91  |  Page 92  |  Page 93  |  Page 94  |  Page 95  |  Page 96  |  Page 97  |  Page 98  |  Page 99  |  Page 100  |  Page 101  |  Page 102  |  Page 103  |  Page 104  |  Page 105  |  Page 106  |  Page 107  |  Page 108  |  Page 109  |  Page 110  |  Page 111  |  Page 112  |  Page 113  |  Page 114  |  Page 115  |  Page 116  |  Page 117  |  Page 118  |  Page 119  |  Page 120  |  Page 121  |  Page 122  |  Page 123  |  Page 124