The advent of animal (metazoan) biomineralization marks a fundamental transition in Earth's history, facilitating the diversification of novel body plans as well as the biological control of carbonate sediment production, resulting in a permanent restructuring of the global carbon cycle. The oldest known skeletal metazoans, the tubular ‘cloudinids’ e.g., Cloudina, appeared during the terminal Ediacaran of the Nama Group, Namibia, but precisely when, where, and why metazoans first acquired the ability to biomineralize and their timing relative to the regional δ13Ccarb profile is undocumented. Assessing possible environmental triggers for this key event therefore requires accurate constraint of the age, paleoenvironmental setting and the geochemical context of the earliest cloudinids.Here we present new stratigraphic, sedimentological, and geochemical (δ13Ccarb and δ18Ocarb) data from the oldest strata (Dabis Formation) of the Nama Group from the Witputs Sub-basin, exposed in the Tsaus Mountains of the Tsau Khaeb National Park (formerly Sperrgebiet). This constrains the oldest confirmed occurrence of Cloudina to limestones of the lower Kliphoek Member, ca. 551–550 Ma, that were deposited laterally-equivalent to the more proximal fossiliferous Kliphoek quartzite. These limestones have dominantly negative δ13Ccarb values that immediately precede recovery from the basal Nama negative δ13Ccarb excursion (BANE), and were deposited in shallow waters after a transition from semi-restricted, evaporitive-dolomitic to open marine carbonate settings. Regional correlation shows that Cloudina first appeared during an interval of dominantly low oxygen and unstable, regional marine redox conditions. We conclude that data do not support a long-term shift towards more stable, oxygenated conditions as a driver for the first appearance of skeletonization in metazoans, but do suggest that open marine carbonate settings with high carbonate supersaturation were required to support the calcifying Cloudina. In the Nama Group, the appearance of Cloudina also notably coincided with a major sea level lowstand, which we hypothesise may have enabled oxygenation of these more distal settings. Such an expansion of shallow marine oxygen may therefore have provided an evolutionary driver for innovations that were not metabolically costly, such as the onset of Cloudina biomineralization.
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