The evolutionary events during the Ediacaran-Cambrian transition (~541 Myr ago) are unparalleled in Earth history. The fossil record suggests that most extant animal phyla appeared in a geologically brief interval, with the oldest unequivocal bilaterian body fossils found in the Early Cambrian. Molecular clocks and biomarkers provide independent estimates for the timing of animal origins, and both suggest a cryptic Neoproterozoic history for Metazoa that extends considerably beyond the Cambrian fossil record. We report an assemblage of ichnofossils from Ediacaran-Cambrian siltstones in Brazil, alongside U-Pb radioisotopic dates that constrain the age of the oldest specimens to 555-542 Myr. X-ray microtomography reveals three-dimensionally preserved traces ranging from 50 to 600 μ m in diameter, indicative of small-bodied, meiofaunal tracemakers. Burrow morphologies suggest they were created by a nematoid-like organism that used undulating locomotion to move through the sediment. This assemblage demonstrates animal-sediment interactions in the latest Ediacaran period, and provides the oldest known fossil evidence for meiofaunal bilaterians. Our discovery highlights meiofaunal ichnofossils as a hitherto unexplored window for tracking animal evolution in deep time, and reveals that both meiofaunal and macrofaunal bilaterians began to explore infaunal niches during the late Ediacaran. NATuRE ECoLoGy & EvoLuTIoN Articles Nature ecology & evolutioNsupport for these suggestions is limited to purported body fossils of sponges 19 and demosponge biomarkers 20 . A considerable gap therefore remains between the fossil record of the late Ediacaran and molecular clock estimates for deep splits in the animal tree, for example the origin of Metazoa and Eumetazoa 3 . Assuming that contemporary molecular clock analyses yield accurate, if imprecise 18 , node ages for animal divergences, a small body size and concomitant limited fossilization potential 21 could reconcile these discordant records of animal evolution (but see ref. 22 ).The small body size of the ancestral bilaterian is supported by recent phylogenomic analyses of deep animal relationships, with acoel flatworms and xenoturbellids (Xenacoelomorpha) being a sister group to all remaining bilaterians (Nephrozoa) 23 , and smallbodied spiralian taxa (the 'Platyzoa') recognized as a paraphyletic grade with respect to macroscopic trochozoans 24 . This suggests that early bilaterians and spiralians were small bodied, possibly meiofaunal, and moved using ciliary gliding.Meiofauna comprises all organisms between 32 and 1,000 μ m in size that inhabit pore-water-rich sediments in freshwater to deepmarine environments 25 . Modern meiofaunal communities include animals, foraminifera and some ciliates, and contribute significantly to sediment bioturbation and bioirrigation 26,27 . The meiofauna can be divided into permanent members (that is, animals with organisms of a small size adapted and restricted to the meiofaunal, interstitial realm) and temporary meiofauna (for example, the ...

Diploblastic eumetazoans of the phylum Cnidaria originated during the Neoproterozoic Era, possibly during the Cryogenian Period. The oldest known fossil cnidarians occur in strata of Ediacaran age and consist of polypoid forms that were either nonbiomineralizing or weakly so. The oldest possible anthozoans, including the genus Ramitubus, may be related to tabulate corals and occur in the Doushantuo Lagerstätte (upper Doushantuo Formation, South China), the age of which is poorly constrained (approximately 585 Ma?). Conulariid scyphozoans may first appear as early as 635–577 Ma (Lantian Formation, South China). A definite conulariid, most similar to Palaeozoic species assigned to the genus Paraconularia, occurs in association with the possible scyphozoan, Corumbella werneri, in the latest Ediacaran (c. 543 Ma) Tamengo Formation of Brazil. Basal Cambrian (c. 540 Ma) phosphorites in the upper Kuanchuanpu Formation (South China) yield solitary polyps of the oldest probable anthozoan (Eolympia pediculata), which appears to have been a stem hexacorallian. This same formation contains fossils interpreted by some authors as pentaradial cubozoan polyps; however, both the oldest known cubozoans and the oldest hydrozoans, all medusae, may actually occur in the Cambrian (Series 3, c. 505 Ma) Marjum Formation (Utah, USA). Although these recently published palaeontological data tend to corroborate the hypothesis that Cnidaria has a relatively deep Neoproterozoic history, the timing of major internal branching events remains poorly constrained, with, for example, the results of some molecular clock analyses indicating that the two cnidarian subphyla (Anthozoaria and Medusozoa) may have originated as many as one billion years ago. Further progress towards elucidating the evolution and early fossil record of cnidarians may accrue from: (1) an intensive search for phosphatized soft parts in possible anthozoans from the Ediacaran Doushantuo Formation; (2) an expanded search for Ediacaran conulariids; and (3) additional detailed analyses of the taphonomy and preservation of Ediacaran and Cambrian cnidarians, including possible pentaradial cubozoan polyps from the Fortunian upper Kuanchuanpu Formation.

The Ediacaran fossil Corumbella is important because it is hypothesized to be a scyphozoan cnidarian, and thus might be one of the rare examples of bona fide Neoproterozoic animals. Unfortunately, its mode of life, style of skeletonization, and taxonomic affinity have been very controversial. Here, we use X-ray micro-CT, SEM, and taphonomic analysis to compare preservational modes of Corumbella, in order to better understand the symmetry, mode of construction, preservational style, and taxonomy of this group. Results suggest that articulated and disarticulated specimens of Corumbella from the Ediacaran of Brazil, Paraguay, and the United States, although sometimes preserved very differently, represent the same taxon—Corumbella werneri. Corumbellids had a thick but flexible theca and probably lived with their basalmost part anchored in the sediment, much like Conotubus. When considered together, these results suggest that Corumbella was one of the first animals to build a skeleton, employing a lamellar microfabric similar to conulariids.

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