Ultrastructure of Ediacaran cloudinids suggests diverse taphonomic histories and affinities with non-biomineralized annelids

Cloudinids have long been considered the earliest biomineralizing metazoans, but their affinities have remained contentious and undetermined. Based on well-preserved ultrastructures of two taxa, we here propose new interpretations regarding both their extent of original biomineralization and their phylogenetic affinity. One of these taxa is a new cloudinid from Mongolia, Zuunia chimidtsereni gen. et sp. nov., which exhibits key characteristics of submicrometric kerogenous lamellae, plastic tube-wall deformation, and tube-wall delamination. Multiple carbonaceous lamellae are also discovered in Cloudina from Namibia and Paraguay, which we interpret to have originated from chitinous or collagenous fabrics. We deduce that these cloudinids were predominantly originally organic (chitinous or collagenous), and postmortem decay and taphonomic mineralization resulted in the formation of aragonite and/or calcite. Further, based on our ultrastructural characterization and other morphological similarities, we suggest that the cloudinids should most parsimoniously be assigned to annelids with originally organic tubes.