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An armoured Cambrian lobopodian from China with arthropod-like appendages

Nature volume 470, pages 526530 (24 February 2011) | Download Citation


Cambrian fossil Lagerstätten preserving soft-bodied organisms have contributed much towards our understanding of metazoan origins1,2,3. Lobopodians are a particularly interesting group that diversified and flourished in the Cambrian seas. Resembling ‘worms with legs’, they have long attracted much attention in that they may have given rise to both Onychophora (velvet worms)4,5,6 and Tardigrada (water bears)7,8, as well as to arthropods in general9,10,11,12. Here we describe Diania cactiformis gen. et sp. nov. as an ‘armoured’ lobopodian from the Chengjiang fossil Lagerstätte (Cambrian Stage 3), Yunnan, southwestern China. Although sharing features with other typical lobopodians, it is remarkable for possessing robust and probably sclerotized appendages, with what appear to be articulated elements. In terms of limb morphology it is therefore closer to the arthropod condition, to our knowledge, than any lobopodian recorded until now. Phylogenetic analysis recovers it in a derived position, close to Arthropoda; thus, it seems to belong to a grade of organization close to the point of becoming a true arthropod. Further, D. cactiformis could imply that arthropodization (sclerotization of the limbs) preceded arthrodization (sclerotization of the body). Comparing our fossils with other lobopodian appendage morphologies—see Kerygmachela9,10, Jianshanopodia13 and Megadictyon12—reinforces the hypothesis that the group as a whole is paraphyletic, with different taxa expressing different grades of arthropodization.

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This work was supported by a Research Scholarship from the Alexander von Humboldt Foundation at the Freie Universität Berlin, the National Science Foundation of China (grants 40802011 and 40830208) and the MOST Special Fund from the State Key Laboratory of Continental Dynamics, Northwest University (to J.L.), and a DFG grant within the Forschergruppe 736 (to H.K. and M.S.). We thank M. G. Chi for the reconstruction, J. Evers and M. Y. Sun for photography and table, and local workers for their strenuous field work.

Author information


  1. Early Life Institute, State Key Laboratory of Continental Dynamics, Department of Geology, Northwest University, Xi’an 710069, China

    • Jianni Liu
    • , Degan Shu
    • , Jian Han
    • , Zhifei Zhang
    •  & Xingliang Zhang
  2. Department of Earth Science, Freie Universität Berlin, D-12249 Berlin, Germany

    • Jianni Liu
    • , Michael Steiner
    •  & Helmut Keupp
  3. Museum für Naturkunde, Leibniz Institute for Research on Evolution and Biodiversity at the Humboldt University Berlin, D-10115 Berlin, Germany

    • Jason A. Dunlop
  4. School of Earth Sciences and Resources, China University of Geosciences, Beijing 100083, China

    • Degan Shu
    •  & Qiang Ou


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J.L. collected most of the fossils, described them and wrote the paper with the other authors; M.S. collected material and was involved in the phylogenetic analysis; J.A.D. contributed to the discussion; H.K. and D.S. were involved in technical aspects of the analysis; Q.O. provided three specimens. J.H. and Z.Z. contributed to fieldwork; X.Z. was involved in the analysis.

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The authors declare no competing financial interests.

Corresponding author

Correspondence to Jianni Liu.

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