•New carbon isotope data capture the final stages of the Shuram/Wonoka excursion.•The transition to positive values postdates the appearance of calcified metazoans.•Anoxic ferruginous deeper waters ...were prevalent throughout this time.•Only mid-ramp settings were fully oxygenated.•Stability of oxygenation controlled the ecology of Ediacaran metazoan communities.
The first appearance of skeletal metazoans in the late Ediacaran (∼550 million years ago; Ma) has been linked to the widespread development of oxygenated oceanic conditions, but a precise spatial and temporal reconstruction of their evolution has not been resolved. Here we consider the evolution of ocean chemistry from ∼550 to ∼541Ma across shelf-to-basin transects in the Zaris and Witputs Sub-Basins of the Nama Group, Namibia. New carbon isotope data capture the final stages of the Shuram/Wonoka deep negative C-isotope excursion, and these are complemented with a reconstruction of water column redox dynamics utilising Fe–S–C systematics and the distribution of skeletal and soft-bodied metazoans. Combined, these inter-basinal datasets provide insight into the potential role of ocean redox chemistry during this pivotal interval of major biological innovation.
The strongly negative δ13C values in the lower parts of the sections reflect both a secular, global change in the C-isotopic composition of Ediacaran seawater, as well as the influence of ‘local’ basinal effects as shown by the most negative δ13C values occurring in the transition from distal to proximal ramp settings. Critical, though, is that the transition to positive δ13C values postdates the appearance of calcified metazoans, indicating that the onset of biomineralization did not occur under post-excursion conditions.
Significantly, we find that anoxic and ferruginous deeper water column conditions were prevalent during and after the transition to positive δ13C that marks the end of the Shuram/Wonoka excursion. Thus, if the C isotope trend reflects the transition to global-scale oxygenation in the aftermath of the oxidation of a large-scale, isotopically light organic carbon pool, it was not sufficient to fully oxygenate the deep ocean.
Both sub-basins reveal highly dynamic redox structures, where shallow, inner ramp settings experienced transient oxygenation. Anoxic conditions were caused either by episodic upwelling of deeper anoxic waters or higher rates of productivity. These settings supported short-lived and monospecific skeletal metazoan communities. By contrast, microbial (thrombolite) reefs, found in deeper inner- and mid-ramp settings, supported more biodiverse communities with complex ecologies and large skeletal metazoans. These long-lived reef communities, as well as Ediacaran soft-bodied biotas, are found particularly within transgressive systems, where oxygenation was persistent. We suggest that a mid-ramp position enabled physical ventilation mechanisms for shallow water column oxygenation to operate during flooding and transgressive sea-level rise. Our data support a prominent role for oxygen, and for stable oxygenated conditions in particular, in controlling both the distribution and ecology of Ediacaran skeletal metazoan communities.
It has been proposed that the terminal Neoproterozoic Ediacara biota were driven to extinction by the evolution of metazoan groups capable of engineering their environments (the ‘biotic replacement’ ...model). However, evidence for an overlapping ecological association between metazoans and soft-bodied Ediacaran organisms is limited. Here, we describe new fossil localities from southern Namibia that preserve soft-bodied Ediacara biota, enigmatic tubular organisms thought to represent metazoans, and vertically-oriented metazoan trace fossils. Although the precise identity of the tracemakers remains elusive, the structures bear several striking similarities with the Cambrian-Recent ichnogenus Conichnus. These new data support inference of stratigraphic and ecological overlap between two very different eukaryotic clades, and indicate the existence of unusual ecosystems comprising both Ediacara biota and metazoans immediately prior to the Cambrian explosion.
•We report the discovery of new fossil sites in the Zaris subbasin, Namibia, preserving soft-bodied Ediacara biota, metazoan body fossils, and metazoan trace fossils•Close stratigraphic relationships among these fossils illustrate that Ediacarans and metazoans occupied the same communities prior to the Cambrian•Spatial relationships suggest niche partitioning between Ediacara biota and metazoan trace makers latest Ediacaran ‘transitional’ ecosystems•Although the nature of biotic interactions between these groups is unclear, these fossils fulfill a prediction of the ‘biotic replacement model’•This supports the notion that the extinction and decline of the Ediacara biota was driven first by metazoan radiation and ecosystem engineering
•Redox sensitive elements in Ediacaran early marine reef cements reveal changes in water column redox.•Marine cement analysis enables exploration of shorter timescales of redox variation than ...standard geochemical techniques.•Ediacaran benthic communities grew close to the chemocline.•Ferruginous, dolomitised sediment and replaced cement may record a transient incursion of anoxic waters.•Communities may have been terminated by dolomitising, anoxic waters that shoaled during short-term, transgressive cycles.
The Ediacaran oceanic redox landscape was heterogeneous, where many basins had a shallow and highly dynamic chemocline above anoxic (ferruginous or euxinic) or low oxygen (manganous) waters. Seawater mMg/Ca ratio was also high, promoting early diagenetic dolomitisation. How the benthos responded to these conditions is fundamental to understanding their ecological dynamics. Here we utilise redox sensitive elements in early marine carbonate cements to investigate possible water column redox controls on the distribution and growth of the oldest metazoan communities.
Skeletal communities in the Zaris Sub-Basin of the Nama Group, Namibia (∼550–547 Ma), grew in shallow waters where fine-grained carbonate sediment often shows evidence of early dolomitisation. Mid-ramp Cloudina reefs are composed of open, highly porous structures that formed multiple, successive assemblages. Each assemblage is terminated by thin (<1 mm), layers of dolomicrite sediment and dolomite cement. All dolomitic lithologies in the Nama Group analysed via Fe speciation suggest precipitation under anoxic ferruginous water column conditions.
Reef cements show a paragenetic sequence from synsedimentary to early marine cement and final burial, which we infer were precipitated under dynamic redox conditions. First, acicular pseudomorphed aragonite cement formed under oxic conditions (low Fe and Mn). Next, the presence of iron-rich dolomicrite sediment, often associated with a recrystallised ferroan dolomite crust, suggests that originally aragonitic or calcitic sediment and a high-Mg precursor cement were preferentially dolomitised. Dolomitisation may have been enhanced either via upwelling of deeper water, anoxic, ferruginous seawater, or by later fluid remobilisation from adjacent shales. A following Mn-rich calcite cement is inferred to be early marine due to its inclusion-rich, fibrous form and well-preserved CL zonation. The final blocky cement precipitated under oxic conditions, probably during shallow burial.
The cements likely record a general shallow to deeper water transect, from oxic shallow waters to low oxygen manganous waters, then to oxic, shallow burial conditions. We hypothesize that Cloudina reef communities were terminated by episodic sediment incursions during short-term, transgressive cycles, possibly accompanied by upwelled, anoxic, ferruginous and dolomitising waters, although the timing for this is poorly constrained. More generally, such incursions may have terminated Ediacaran benthic communities that grew close to the chemocline.
•δ44Ca from limestones aged ∼550–539 Ma that host the earliest skeletal animal fossils.•Negative shift in δ44Ca that lasted at least 11–14Myrs.•Unlikely to record a transition towards more ...sediment-buffered carbonate diagenesis.•May record enhanced continental weathering or evaporite deposition.•May record a change in timing of dolomitisation.
Calcium isotope ratios in ancient carbonate rocks can provide insight into the global marine calcium cycle as well as local conditions during carbonate mineral precipitation and diagenesis. We compare two extraction techniques for the separation of calcium from other ions before δ44Ca analysis, using an automated ion chromatograph and using manual gravity columns. The two techniques produce the same δ44Ca within error (2σ). We present 31 δ44Ca analyses of carbonate rocks from the Nama Group, Namibia, which record a negative shift in δ44Ca of 0.35‰ between ∼550 and ∼547 Ma, from −1.25‰ to −1.60‰, followed by persistently low δ44Ca (−1.48±0.06‰) between ∼547 and 539 Ma. Very low δ44Ca (<−1.5‰) are commonly interpreted to represent the preservation of local aragonite that has recrystallized to calcite under sediment-buffered conditions (where the composition of the diagenetic carbonate product is determined mainly by the original sediments). The shift in δ44Ca across the Nama Group could therefore represent a change from fluid-buffered diagenesis (where the composition of the diagenetic carbonate mineral is determined mainly by the fluid) to sediment-buffered diagenesis. However, this interpretation is not consistent with either potential geochemical indicators of diagenesis (e.g., δ18O), or changes in large-scale fluid-flow as predicted from sequence stratigraphy. We consider alternative interpretations for generating changes in the δ44Ca of ancient carbonate rocks including enhanced continental weathering, increases in evaporite deposition, and changes in the style of dolomitisation.
•A new late Neoproterozoic sequence in of Southern Africa.•Drivers for Ediacaran evolution just prior to their great extinction.•Geochemistry/geochronology of late Neoproterozoic Nama Group ...paleoenvironment.
Common, Ediacaran fossils are well preserved in a Late Neoproterozoic (ca. 545Ma) shallow marine sequence, described here as the Aar Member of the Dabis Formation (Kuibis Subgroup, Nama Group), near Aus in southwest Namibia. This 31–38m thick, shale-dominant unit records the transition from fluvial-shallow marine Kliphoek Sandstone to open marine limestone of the Mooifontein Member of the Zaris Formation, deposited on a subsiding continental margin during a major, regional transgression. Thin sandstone beds contain fossils at a number of levels throughout the Aar Member. Concentrations of Pteridinium were mostly transported in flood-derived sheets, while some Ernietta assemblages are preserved close to in situ. Rangea has also been transported, and is mostly confined to thin sandstone lenses incised into mudstone. Limestone beds, common throughout, include at least two marker horizons that can be followed regionally and show local evidence of storm reworking. Systematic sampling and analyses of limestone reveals enrichment in both 13C and 18O higher in the section, with negative δ13C near the base rising to moderate positive values near the top. The negative-to-positive transition in δ13C values is more pronounced in the east, with all of the lower Aar Member samples consistently depleted in 13C. While this may reflect greater degrees of alteration by meteoric or dewatering fluids, the same carbonates are notably enriched in 18O relative to those at the same stratigraphic position to the west. The overall rise in 13C is attributed to greater proportional burial of organic matter and release of oxygen to surface environments, while the spatial variability is likely the result of a strong surface-to-deep carbon isotopic gradient in seawater. A number of the fossils, especially Rangea, are encrusted with jarosite, an iron-bearing sulphate mineral and common weathering product of pyrite. This observation suggests that preservation of the fossils may have resulted from the rapid encrustation of pyrite on the surface of the organisms as they decomposed and were consumed by sulphate-reducing bacteria within the sandy, near shore sediments. Insofar as pyrite formation requires iron, which is soluble and reactive in anoxic solutions, it is likely that the deeper subtidal environments lacked oxygen. In situ pyritized forms like Ernietta may have developed the capacity to survive under episodically anoxic or sub-oxic environmental conditions, while Pteridinium and Rangea lived within an oxygenated estuarine or fluvial setting and were transported during storms to anoxic, ferruginous environments where they were exquisitely preserved.
Here we describe large, complex trace fossils in the late Ediacaran Omkyk Member of the Zaris Formation, Nama Group, southern Namibia. The horizontal trace fossils are preserved on a number of talus ...blocks from a bedding plane of a cm-thick sandstone lens from a single stratigraphic horizon less than 100 m below an ash bed dated at 547.3 ± 0.7 Ma. The forms consist of overlapping U-shaped spreiten elements with parallel limbs surrounded by an outer tube. Individual U-shaped elements are 0.2 to 1 cm in diameter, the outer tube is less than 3 mm in diameter, and the forms as a whole range from 5 to 30 cm long and 3 to 10 cm wide. The specimens commonly show a change in direction and change in diameter. The morphology of these trace fossils is comparable to backfill structures, particularly specimens of Paleozoic Zoophycos from shallow water environments. Here we interpret these horizontal spreiten-burrows to record the grazing of the trace-maker on or below a textured organic surface. The identification of large late Ediacaran trace fossils is consistent with recent reports of backfilled horizontal burrows below the Precambrian–Cambrian boundary and is suggestive of the appearance of complex feeding habits prior to the Cambrian trace fossil explosion.
Reefs containing abundant calcified metazoans occur at several stratigraphic levels within carbonate platforms of the terminal Proterozoic Nama Group, central and southern Namibia. The reef-bearing ...strata span an interval ranging from approximately 550 Ma to 543 Ma. The reefs are composed of thrombolites (clotted internal texture) and stromatolites (laminated internal texture) that form laterally continuous biostromes, isolated patch reefs, and isolated pinnacle reefs ranging in scale from a meter to several kilometers in width. Stromatolite-dominated reefs occur in depositionally updip positions within carbonate ramps, whereas thrombolite-dominated reefs occur broadly across the ramp profile and are well developed as pinnacle reefs in downdip positions. The three-dimensional morphology of reef-associated fossils was reconstructed by computer, based on digitized images of sections taken at 25-micron intervals through 15 fossil specimens and additionally supported by observations of over 90 sets of serial sections. Most variation observed in outcrop can be accounted for by a single species of cm-scale, lightly calcified goblet-shaped fossils herein described as Namacalathus hermanastes gen. et sp. nov. These fossils are characterized by a hollow stem open at both ends attached to a broadly spheroidal cup marked by a circular opening with a downturned lip and six (or seven) side holes interpreted as diagenetic features of underlying biological structure. The goblets lived atop the rough topography created by ecologically complex microbial-algal carpets; they appear to have been sessile benthos attached either to the biohermal substrate or to soft-bodied macrobenthos such as seaweeds that grew on the reef surface. The phylogenetic affinities of Namacalathus are uncertain, although preserved morphology is consistent with a cnidarian-like bodyplan. In general aspect, these fossils resemble some of the unmineralized, radially symmetric taxa found in contemporaneous sandstones and shales, but do not appear to be closely related to the well-skeletonized bilaterian animals that radiated in younger oceans. Nama reefs demonstrate that biohermal associations of invertebrates and thrombolite-forming microorganisms antedate the Cambrian Period.
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