Macroscopic carbonaceous compressions in a terminal Proterozoic shale; a systematic reassessment of the Miaohe biota, South China

Carbonaceous compression fossils in shales of the uppermost Doushantuo Formation (ca. 555-590 Ma) at Miaohe in the Yangtze Gorges area provide a rare Burgess-Shale-type taphonomic window on terminal Proterozoic biology. More than 100 macrofossil species have been described from Miaohe shales, but in an examination of published and new materials, we recognize only about twenty distinct taxa, including Aggregatosphaera miaoheensis new gen. and sp. Most of these fossils can be interpreted unambiguously as colonial prokaryotes or multicellular algae. Phylogenetically derived coenocytic green algae appear to be present, as do regularly bifurcating thalli comparable to red and brown algae. At least five species have been interpreted as metazoans by previous workers. Of these, Protoconites minor and Calyptrina striata most closely resemble animal remains; either or both could be the organic sheaths of cnidarian scyphopolyps, although an algal origin cannot be ruled out for P. minor. Despite exceptional preservation, the Miaohe assemblage contains no macroscopic fossils that can be interpreted with confidence as bilaterian animals. In combination with other late Neoproterozoic and Early Cambrian body fossils and trace fossils, the Doushantuo assemblage supports the view that body-plan diversification within bilaterian phyla was largely a Cambrian event.