The rise of multicellularity represents a major evolutionary transition and it occurred independently in multiple eukaryote clades. Although simple multicellular organisms may have evolved in the Mesoproterozoic Era or even earlier, complex multicellular eukaryotes began to diversify only in the Ediacaran Period, just before the Cambrian explosion. Thus, the Ediacaran fossil record can provide key paleontological evidence about the early radiation of multicellular eukaryotes that ultimately culminated in the Cambrian explosion. The Ediacaran Weng'an biota in South China hosts exceptionally preserved eukaryote fossils, including various acanthomorphic acritarchs, pseudoparenchymatous thalli, tubular microfossils, and spheroidal fossils such as Megasphaera, Helicoforamina, Spiralicellula, and Caveasphaera. Many of these fossils have been interpreted as multicellular eukaryotes, although alternative interpretations have also been proposed. In this review, we critically examine these various interpretations, focusing particularly on Megasphaera, which has been variously interpreted as a sulfur-oxidizing bacterium, a unicellular protist, a mesomycetozoean-like holozoan, a volvocine green alga, a stem-group animal, or a crown-group animal. We conclude that Megasphaera is a multicellular eukaryote with evidence for cell-to-cell adhesion, a flexible membrane unconstrained by a rigid cell wall, spatial cellular differentiation, germ–soma separation, and programmed cell death. These features are inconsistent with the bacterium, unicellular protist, and mesomycetozoean-like holozoan interpretations. Thus, the surviving hypotheses, particularly the stem-group animal and algal interpretations, should be further tested with additional evidence. The Weng'an biota also hosts cellularly differentiated pseudoparenchymatous thalli with specialized reproductive structures indicative of an affinity with florideophyte red algae. The other Weng'an fossils reviewed here may also be multicellular eukaryotes, although direct cellular evidence is lacking in some and phylogenetic affinities are poorly constrained in others. The Weng'an biota offers many research opportunities to resolve the life histories and phylogenetic diversity of early multicellular eukaryotes and to illuminate the evolutionary prelude to the Cambrian explosion.
Rocks of Ediacaran age (~635–541 Ma) contain the oldest fossils of large, complex organisms and their behaviors. These fossils document developmental and ecological innovations, and suggest that extinctions helped to shape the trajectory of early animal evolution. Conventional methods divide Ediacaran macrofossil localities into taxonomically distinct clusters, which may represent evolutionary, environmental, or preservational variation. Here, we investigate these possibilities with network analysis of body and trace fossil occurrences. By partitioning multipartite networks of taxa, paleoenvironments, and geologic formations into community units, we distinguish between biostratigraphic zones and paleoenvironmentally restricted biotopes, and provide empirically robust and statistically significant evidence for a global, cosmopolitan assemblage unique to terminal Ediacaran strata. The assemblage is taxonomically depauperate but includes fossils of recognizable eumetazoans, which lived between two episodes of biotic turnover. These turnover events were the first major extinctions of complex life and paved the way for the Cambrian radiation of animals.
In terminal Ediacaran strata of South China, the onset of calcareous biomineralization is preserved in the paleontological transition from Conotubus to Cloudina in repetitious limestone facies of the Dengying Formation. Both fossils have similar size, funnel-in-funnel construction, and epibenthic lifestyle, but Cloudina is biomineralized, whereas Conotubus is not. To provide environmental context for this evolutionary milestone, we conducted a high-resolution elemental and stable isotope study of the richly fossiliferous Gaojiashan Member. Coincident with the first appearance of Cloudina is a significant positive carbonate carbon isotope excursion (up to +6‰) and an increase in the abundance and (34) S composition of pyrite. In contrast, δ(34) S values of carbonate-associated sulfate remain steady throughout the succession, resulting in anomalously large (>70‰) sulfur isotope fractionations in the lower half of the member. The fractionation trend likely relates to changes in microbial communities, with sulfur disproportionation involved in the lower interval, whereas microbial sulfate reduction was the principal metabolic pathway in the upper. We speculate that the coupled paleontological and biogeochemical anomalies may have coincided with an increase in terrestrial weathering fluxes of sulfate, alkalinity, and nutrients to the depositional basin, which stimulated primary productivity, the spread of an oxygen minimum zone, and the development of euxinic conditions in subtidal and basinal environments. Enhanced production and burial of organic matter is thus directly connected to the carbon isotope anomaly, and likely promoted pyritization as the main taphonomic pathway for Conotubus and other soft-bodied Ediacara biotas. Our studies suggest that the Ediacaran confluence of ecological pressures from predation and environmental pressures from an increase in seawater alkalinity set the stage for an unprecedented geobiological response: the evolutionary novelty of animal biomineralization.
Laflamme, M., Schiffbauer, J.D., Narbonne, G.M., & Briggs, D.E.G. 2011: Microbial biofilms and the preservation of the Ediacara biota. Lethaia, Vol. 44, pp. 203–213. The terminal Neoproterozoic Ediacaran Period is typified by the Ediacara biota (ca. 579–542 Ma), which includes the first morphologically complex macroscopic organisms. Both the taphonomic setting that promoted the preservation of the soft‐bodied Ediacara biota in coarse‐grained sediments, and the influence of associated microbial coatings on this process, have generated debate. Specimens of Ediacaran discs (Aspidella) from the Fermeuse Formation of Newfoundland, Canada, were analysed using environmental scanning electron microscopy (ESEM) and focused ion beam electron microscopy (FIB‐EM) to determine the relationship between the fossil specimens and the surrounding sediment. The presence of chemically distinct (Al–Mg–Fe–K‐ and to a lesser extent S‐rich), finer‐grained sediment (with organized iron sulphides) surrounding the upper and lower margins of the Ediacaran fossils is consistent with elemental analyses of well preserved bacterial biofilms from other localities. ESEM analyses reveal a contrast in the composition of the sediment bound within the discs, which contains a higher concentration of Al, Ca and K, and the purer Si‐rich sediment that forms the surrounding matrix. This suggests that the coarse grained sediment was incorporated into the organism during life. Ediacaran discs were likely surrounded by a bacterial biofilm or thin microbial mat composed primarily of extracellular polymeric substances (or exopolysaccharide) during life, which added structural stability to these frond holdfasts, and facilitated their fossilization. Microbially mediated preservation in Fermeuse‐style Ediacaran taphonomy provides an explanation for the dominance of Aspidella holdfasts in these settings, and suggests that preservation of Ediacaran fossils in the round may be much more prevalent than previously recognized. We suggest that the overwhelming dominance of circular to bulbous forms such as Aspidella in Ediacaran biotas around the world is a direct result of the interplay between microbial ecology and microbially mediated taphonomy. □Aspidella, Ediacaran preservation, environmental scanning electron microscopy, focused ion beam electron microscopy, palaeoecology, taphonomic bias.
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