Meroblastic cleavage identifies some Ediacaran Doushantuo (China) embryo-like fossils as metazoans

Ediacaran Doushantuo (Formation) embryo-like fossils (EDEFs, ca. 600 Ma) from South China display cellular and sub-cellular structures and provide a unique window on the early evolution of multicellular eukaryotes. But there have been widely disparate interpretations of these fossils. Here we report new fossil embryo-like forms from the Doushantuo phosphorite that exhibit a meroblastic cleavage pattern. Our results from high-resolution propagation phase contrast-synchrotron radiation X-ray microtomography (PPC-SRmu CT) demonstrate that these fossils preserve features directly comparable to those of modern meroblastic animal embryos that utilize discoidal cleavage. Given that discoidal-type meroblastic cleavage occurs only in metazoans, the phylogenetic positions of these fossils probably fall into the animal branch of the holozoan tree. Meroblastic as well as holoblastic cleavage forms were thus present by ca. 600 Ma, substantiating the conclusion derived from molecular clock estimates that a variety of metazoan lineages had evolved by the mid-Ediacaran after the termination of the Marinoan glaciation, if not earlier.