A Tremadocian (Early Ordovician) palaeoscolecidan worm from graptolitic shales in Hunan Province, South China

57 3 May 657 671 10.1111/pala.12083

WANG, W., MUIR, L.A., BOTTING, J.P., FENG, H., SERVAIS, T., LI, L., 2014, A Tremadocian (Early Ordovician) palaeoscolecidan worm from graptolitic shales in Hunan Province, South China. Palaeontology, 57, 3, 657–671. doi: 10.1111/pala.12083

<p>Wenhui Wang, Lucy A. Muir, Joseph P. Botting, Hongzhen Feng, Thomas Servais and Lixia Li</p>
  • Wenhui Wang - School of Earth Sciences and Engineering, Nanjing University, Nanjing, China (email: wwhever@126.com)
  • Wenhui Wang - State Key Laboratory of Palaeobiology and Stratigraphy, Key Laboratory of Economic Stratigraphy and Palaeogeography, Nanjing Institute of Geology and Palaeontology, Nanjing, China
  • Lucy A. Muir - State Key Laboratory of Palaeobiology and Stratigraphy, Key Laboratory of Economic Stratigraphy and Palaeogeography, Nanjing Institute of Geology and Palaeontology, Nanjing, China (email: ucy@asoldasthehills.org)
  • Joseph P. Botting - State Key Laboratory of Palaeobiology and Stratigraphy, Key Laboratory of Economic Stratigraphy and Palaeogeography, Nanjing Institute of Geology and Palaeontology, Nanjing, China (email: acutipuerilis@yahoo.co.uk)
  • Hongzhen Feng, - School of Earth Sciences and Engineering, Nanjing University, Nanjing, China (email: feng.h.zh@163.com)
  • Thomas Servais - Géosystèmes, UMR 8217 du CNRS, Université Lille 1, Avenue Paul Langevin, Villeneuve d'Ascq, France (email: thomas.servais@univ-lille1.fr)
  • Lixia Li - School of Earth Sciences and Engineering, Nanjing University, Nanjing, China (email: lilixia1015@gmail.com)
National Natural Science Foundation of China. Grant Numbers: 41230208, 41372017, 41172012 CNRS, Universities of Lyon1 and Lille1

A new locality exposing Tremadocian (Early Ordovician) graptolitic shales in Hunan Province, South China, has yielded an exceptionally well-preserved annulated worm. This palaeoscolecidan is described as Waflascolex changdensis gen. et sp. nov. and reveals extremely fine detail of the cuticle organization and plating array. The new taxon is characterized by three critical characters: incomplete plate rows that occur only on the posterior end of the worm (except in the posterior-most area) and do not extend over the entire circumference or along the entire trunk; a regular rhomboidal array of platelets around intercalations; and reduced cuticular organization at the posterior termination. The unique cuticular organization and platelet ornamentation in the new taxon offer insight into functional differentiation of plates in the scleritome. Palaeoscolecid distribution through the early Palaeozoic is reviewed, showing that the worms were widespread in the Cambrian and Ordovician, but became more restricted during the Silurian. Ordovician palaeoscolecidans are diverse in scleritome architecture, and strikingly different from Cambrian taxa, indicating that this group diversified as part of the Great Ordovician Biodiversification Event.

  • Bather, F. A. 1920. Protoscolex latus, a new “worm” from Lower Ludlow beds. Annals and Magazine of Natural History, Series 9, 5, 124–133.
  • Bendix-Almgreen, S. E. and Peel, J. S. 1988. Hadimopanella from the Lower Cambrian of North Greenland: structure and affinities. Bulletin of the Geological Society of Denmark, 37, 83–103.
  • Boogard, M. van den 1988. Some data on Milaculum Müller, 1973. Scripta Geologica, 88, 1–25.
  • van den Boogard, M. 1989. Isolated tubercules of some Palaeoscolecida. Scripta Geologica, 90, 1–12.
  • Botting, J. P. and Muir, L. A. 2012. Fauna and ecology of the Holothurian Bed, Llandrindod, Wales, UK (Darriwilian, Middle Ordovician), and the oldest articulated holothurian. Palaeontologia Electronica, 15 (1) 9A, 28 pp.
  • Botting, J. P., Muir, L. A., Van Roy, P., Bates, D. and Upton, C. 2012. Diverse Middle Ordovician palaeoscolecidan worms from the Builth-Llandrindod Inlier of central Wales. Palaeontology, 55, 501–528.
  • Brock, G. A. and Cooper, B. J. 1993. Shelly fossils from the Early Cambrian (Toyonian) Wirrealpa, Aroona Creek, and Ramsay limestones of South Australia. Journal of Paleontology, 67, 758–787.
  • Bulman, O. M. B. 1927. A monograph of British dendroid graptolites, part 1. Palaeontographical Society Monograph, 79, 1–28.
  • Chen, X., Rong, J. Y., Wang, X. F., Wang, Z. H., Zhang, Y. D. and Zhan, R. B. 1995. Correlation of the Ordovician rocks of China: charts and explanatory notes, Vol. 31. International Union of Geological Sciences Publication, Norway, 104 pp.
  • Cisne, J. L. 1973. Beecher's Trilobite Bed revisited: ecology of an Ordovician deepwater fauna. Postilla, 160, 1–25.
  • Clark, D. L., Sorenson, J. K., Ladd, A. N. and Freiheit, J. R. 1999. Probable microvertebrates, vertebrate-like fossils, and weird things from the Wisconsin Ordovician. Journal of Paleontology, 73, 1201–1209.
  • Conway Morris, S. 1977. Fossil priapulid worms. Special Papers in Palaeontology, 20, 103 pp.
  • Conway Morris, S. 1997. The cuticular structure of the 495-Myr-old type species of the fossil worm Palaeoscolex, P. piscatorum (?Priapulida). Zoological Journal of the Linnaean Society, 119, 69–82.
  • Conway Morris, S., Peel, J. S. 2010. New palaeoscolecidan worms from the lower Cambrian: Sirius Passet Fossil-Lagersttte (North Greenland), Latham Shale (California), and Kinzers Shale (Pennsylvania). Acta Palaeontologica Polonica, 51, 141–156.
  • Conway Morris, S. and Robison, R. A. 1986. Middle Cambrian priapulids and other soft-bodied fossils from Utah and Spain. The University of Kansas Paleontological Contributions, 117, 22 pp.
  • Conway Morris, S., Pickerill, R. K. and Harland, T. L. 1982. A possible annelid from the Trenton Limestone (Ordovician) of Quebec, with a review of fossil oligochaetes and other annulate worms. Canadian Journal of Earth Sciences, 19, 2150–2157.
  • Cooper, R. A., Maletz, J., Wang, H. F. and Erdtmann, B. D. 1998. Taxonomy and evolution of earliest Ordovician graptoloids. Norsk Geologisk Tidsskrift, 78, 3–32.
  • Dornbos, S. Q. and Chen, J. Y. 2008. Community palaeoecology of the early Cambrian Maotianshan Shale biota: ecological dominance of priapulid worms. Palaeogeography, Palaeoclimatology, Palaeoecology, 258, 200–212.
  • 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, 55, 613–622.
  • García-Bellido, D. C. and Aceñolaza, G. F. 2011. The worm Palaeoscolex from the Cambrian of NW Argentina: extending the biogeography of Cambrian priapulids to South America. Alcheringa, 35, 531–538.
  • García-Bellido, D. C., Gutiérrez-Marco, J. C. and Chacaltana, C. A. 2008. First soft-bodied fossil from the Ordovician of Peru. Alcheringa, 32, 313–320.
  • García-Bellido, D. C., Paterson, J. R. and Edgecombe, G. D. 2013. Cambrian palaeoscolecids (Cycloneuralia) from Gondwana and reappraisal of species assigned to Palaeoscolex. Gondwana Research, 24, 780–795.
  • Gedik, I. 1977. Conodont biostratigraphy in the Middle Taurus. Bulletin of the Geological Society of Turkey, 20, 34–48.
  • Gedik, I. 1989. Hadimopanellid biostratigraphy in the Cambrian of the Western Taurids: a new biostratigraphic tool in the subdivision of Cambrian System. Geological Bulletin of Turkey, 32, 65–78.
  • Glaessner, M. F. 1979. Lower Cambrian Crustacea and annelid worms from Kangaroo Island, South Australia. Alcheringa, 3, 21–31.
  • Han, J., Zhang, X. L., Zhang, Z. F. and Shu, D. G. 2003. A new platy-armored worm from the Early Cambrian Chengjiang Lagersttte, South China. Acta Geologica Sinica, 77, 1–6.
  • Han, J., Zhang, Z. F. and Liu, J. N. 2008. A preliminary note on the dispersal of the Cambrian Burgess Shale-type faunas. Gondwana Research, 14, 269–276.
  • Harvey, T. H. P., Dong, X. and Donoghue, P. C. J. 2010. Are palaeoscolecids ancestral ecdysozoans? Evolution and Development, 12, 177–200.
  • Hinz, I., Kraft, P., Mergl, M. and Müller, K. J. 1990. The problematic Hadimopanella, Kaimenella, Milaculum and Utahphospha identified as sclerites of Palaeoscolecida. Lethaia, 23, 217–221.
  • Hou, X. G. and Bergström, J. 1994. Palaeoscolecid worms may be nematomorphs rather than annelids. Lethaia, 27, 11–17.
  • Hou, X. G. and Sun, W. G. 1988. Discovery of Chengjiang Fauna at Meishucun, Jinning, Yunnan. Acta Palaeontologica Sinica, 27, 1–12. [In Chinese].
  • Hu, S. X., Luo, H. L., Hou, S. G. and Erdtmann, B. D. 2007. Eocrinoid echinoderms from the Lower Cambrian Guanshan Fauna in Wuding, Yunnan, China. Chinese Science Bulletin, 52, 717–719.
  • Hu, S. X., Li, Y., Luo, H. L., Fu, X. P., You, T., Pang, J. Y., Liu, Q. and Steiner, M. 2008. New Record of Palaeoscolecids from the Early Cambrian of Yunnan, China. Acta Geologica Sinica, 82, 244–248.
  • Huang, D. Y., Vannier, J. and Chen, J. Y. 2004. Recent Priapulidae and their Early Cambrian ancestors: comparisons and evolutionary significance. Geobios, 37, 217–228.
  • Huang, D. Y., Chen, J. Y. and Vannier, J. 2006. Discussion on the systematic position of the Early Cambrian priapulomorph worms. Chinese Science Bulletin, 51, 243–249.
  • Ivantsov, A. Yu. and Wrona, R. 2004. Articulated palaeoscolecid sclerite arrays from the Lower Cambrian of eastern Siberia. Acta Geologica Polonica, 54, 1–22.
  • Ivantsov, A. Yu., Zhuravlev, A. Yu., Krasilov, V. A., Leguta, A. V., Mel'nikova, L. M., Urbanek, A., Ushatinskaya, G. T. and Malakhovskaya, Ya. E. 2005. Unikal'nye sinskie mestonakhozhdeniya rannekembriiskikh organizmov, sibirskaya platforma. [The unique Sinsk locality of Early Cambrian organisms, Siberian Platform.] Trudy paleontologicheskogo instituta, 284, 1–141. [In Russian, English summary].
  • Kraft, P. and Mergl, M. 1989. Worm-like fossils (Palaeoscolecida; ?Chaetognatha) from the Lower Ordovician of Bohemia. Sborník Geologických V&#x11b;d, Paleontologie, 30, 9–36.
  • Lai, C. G. 1982. The Ordovician system of China. Geological Publishing House, Beijing, 294 pp. [In Chinese].
  • Lehnert, O. and Kraft, P. 2006. Manitouscolex, a new palaeoscolecidan genus from the Lower Ordovician of Colorado. Journal of Paleontology, 80, 386–391.
  • Li, Z. X. and Powell, C. McA. 2001. An outline of the paleogeographic evolution of the Australasian region since the beginning of the Neoproterozoic. Earth-Science Reviews, 53, 237–277.
  • Lin, T. R. 1995. Discovery of late Early Cambrian worm from Huainan, Anhui. Acta Palaeontologica Sinica, 34, 505–508.
  • Liu, C. L. 1987. Dendroids from the Strata near the Cambrian-Ordovician Boundary of Taoyuan, Hunan Province. Acta Scientiarum Naturalium Universitatis Sunyatseni, 4, 49–58.
  • LoDuca, S. T. and Brett, C. E. 1997. The Medusaegraptus epibole and Ludlovian Konservat-Lagersttten of eastern North America. 369–405. In Brett, C. E. and Baird, G. (eds). Paleontological events: stratigraphic, ecological and evolutionary implications. Columbia University Press, New York, 604 pp.
  • Luo, H. L., Fu, X. P., Hu, S. X., Li, Y., Hou, S. G., You, T., Pang, J. Y. and Liu, Q. 2007. A new arthropod, Guangweicaris Luo, Fu et Hu gen. nov. from the Early Cambrian Guanshan fauna, Kunming, China. Journal of the Geological Society of China, 81, 1–7.
  • Mikulic, D. G., Briggs, D. E. G. and Klussendorf, J. 1985. A new exceptionally preserved biota from the Lower Silurian of Wisconsin, U.S.A. Philosophical Transactions of the Royal Society of London, Series B: Biological Sciences, 311, 75–85.
  • Miller, S. A. and Faber, C. 1892. Some new species and new structural parts of fossils. Journal of the Cincinnati Society of Natural History, 15, 75–100.
  • Muir, L. A., Ng, T.-W., Li, X.-F., Zhang, Y.-D. and Lin, J.-P. in press. Palaeoscolecidan worms and a possible nematode from the Early Ordovician of South China. Palaeoworld.
  • Müller, K. J. 1973. Milaculum n.g., ein phosphatisches Mikrofossil aus dem Altpalozoikum. Palontologische Zeitschrift, 47, 217–228.
  • Müller, K. J. and Hinz-Schallreuter, I. 1993. Palaeoscolecid worms from the Middle Cambrian of Australia. Palaeontology, 36, 543–592.
  • Owens, R. M., Fortey, R. A., Cope, J. C. W., Rushton, A. W. A. and Bassett, M. G. 1982. Tremadoc faunas from the Carmarthen district, South Wales. Geological Magazine, 119, 1–38.
  • Peng, S. C. 1984. Cambrian-Ordovician boundary in the Cili-Taoyuan border area, northwestern Hunan with description of relative trilobites. 285–405. In Nanjing Institute of Geology and Palaeontology, Academia Sinica (ed.). Stratigraphy and palaeontology of systemic boundaries in China, Cambrian–Ordovician boundary (1). Anhui Science and Technology Publishing House, Hefei, 405 pp.
  • Peng, S. C. 1987. Early Cambrian stratigraphy and trilobite fauna of Taoyuan and Cili, Hunan. 35–134. In Nanjing Institute of Geology and Palaeontology, Academia Sinica (ed.). Collection of postgraduate theses. Nanjing Institute of Geology and Palaeontology, Academia Sinica, Jiangsu Science and Technology Publishing House, Nanjing, 411 pp.
  • Peng, J. 2007. The Qiandongian (Cambrian) Balang Fauna from Eastern Guizhou, South China. Unpublished PhD thesis, Nanjing University, 149 pp.
  • Peng, J., Zhao, Y. L., Wu, Y. S., Yuan, J. L. and Tai, T. S. 2005a. Balang Fauna- A new Early Cambrian fauna from Kaili City, Guizhou. Chinese Science Bulletin, 50, 1159–1162.
  • Peng, J., Zhao, Y. L., Wu, Y. S., Yuan, J. L. and Tai, T. S. 2005b. The discovery of the Early Cambrian Balang fauna in Kaili, Guizhou. Chinese Science Bulletin, 50, 1055–1057. [In Chinese].
  • Repetski, J. E. 1981. An Ordovician occurrence of Utahphospha Müller & Miller. Journal of Paleontology, 55, 395–400.
  • Robison, R. A. 1969. Annelids from the Middle Cambrian Spence Shale of Utah. Journal of Paleontology, 43, 1169–1173.
  • Sadler, P. M., Cooper, R. A. and Melchin, M. 2009. High-resolution, early Paleozoic (Ordovician – Silurian) time scales. Geological Society of America Bulletin, 121, 887–906.
  • Servais, T., Owen, A. W., Harper, D. A. T., Kröger, B. and Munnecke, A. 2010. The Great Ordovician Biodiversification Event (GOBE): the palaeoecological dimension. Palaeogeography, Palaeoclimatology, Palaeoecology, 294, 99–119.
  • Topper, T. P., Brock, G. A., Skovsted, C. B. and Paterson, J. R. 2010. Palaeoscolecid scleritome fragments with Hadimopanella plates from the early Cambrian of South Australia. Geological Magazine, 147, 86–97.
  • Torsvik, T. H. 2009. BugPlates: Linking biogeography and palaeogeography. http://www.geodynamics.no/Web/Content/Software/ (accessed 5 June 2010).
  • Van Roy, P., Orr, P. J., Botting, J. P., Muir, L. A., Vinther, J., Lefebvre, B., el Hariri, K. and Briggs, D. E. G. 2010. Ordovician faunas of Burgess Shale type. Nature, 465, 215–218.
  • Wang, C. Y. 1990. Some Llandovery phosphatic microfossils from South China. Acta Palaeontologica Sinica, 29, 548–556. [In Chinese with English summary].
  • Wang, X. F. and Wang, C. S. 2001. Tremadocian (Ordovician) graptolite diversification events in China. Alcheringa, 25, 155–168.
  • Whittard, W. F. 1953. Palaeoscolex piscatorum gen. et sp. nov., a worm from the Tremadocian of Shropshire. Quarterly Journal of the Geological Society of London, 109, 125–135.
  • Wrona, R. 2004. Cambrian microfossils from glacial erratics of King George Island, Antarctica. Acta Palaeontologica Polonica, 49, 13–56.
  • Yuan, W. W., Zhou, Z. Y., Zhang, J. M., Zhou, Z. Q., Sun, X. W. and Zhou, T. M. 2000. Tremadocian trilobite biofacies in Western Hunan-Hubei. Journal of Stratigraphy, 24, 275–282. [In Chinese].
  • Zhang, X. L. and Hua, H. 2005. Soft-bodied fossils from the Shipai Formation, Lower Cambrian of the Three Gorge area, South China. Geological Magazine, 142, 1–11.
  • Zhang, X. G. and Pratt, B. R. 1996. Early Cambrian Palaeoscolecid cuticles from Shaanxi, China. Journal of Paleontology, 70, 275–279.
  • Zhao, Y. L., Yuan, J. L., Zhu, M. Y., Yang, R. D., Guo, Q. J., Peng, J. and Yang, X. L. 2002. Progress and Significance in Research on the early Middle Cambrian Kaili Biota, Guizhou Province, China. Progress in Natural Science, 12, 649–654.
  • Zhao, Y. L., Zhu, M. Y., Babcock, L. E. and Peng, J. 2011. The Kaili Biota: marine organisms from 508 million years ago. Guizhou Publishing Group, Guiyang, 251 pp. [In Chinese, English figure captions].
Wiley Online Library