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For almost 150 years, megascopic structures in siliciclastic sequences of terminal Precambrian age have been frustratingly difficult to characterize and classify. As with all other areas of human knowledge, progress with exploration, documentation and understanding is growing at an exponential rate. Nevertheless, there is much to be learned from following the evolution of the logic behind the biological interpretations of these enigmatic fossils. Here, I review the history of discovery as well as some long-established core members of widely recognized clades that are still difficult to graft on to the tree of life. These ‘orphan plesions’ occupy roles that were once held by famous former Problematica, such as archaeocyaths, graptolites and rudist bivalves. In some of those cases, taxonomic enlightenment was brought about by the discovery of new characters; in others it required a better knowledge of their living counterparts. Can we use these approaches to rescue the Ediacaran orphans? Five taxa that are examined in this context are Arborea (Arboreomorpha), Dickinsonia (Dickinsoniomorpha), Pteridinium plus Ernietta (Erniettomorpha) and Kimberella (Bilateria?). With the possible exception of Dickinsonia, all of these organisms may be coelenterate grade eumetazoans.

Pteridinium simplex is an iconic erniettomorph taxon best known from late Ediacaran successions in South Australia, Russia, and Namibia. Despite nearly 100 years of study, there remain fundamental questions surrounding the paleobiology and paleoecology of this organism, including its life position relative to the sediment–water interface, and how it fed and functioned within benthic communities. Here, we combine a redescription of specimens housed at the Senckenberg Forschungsinstitut und Naturmuseum Frankfurt with field observations of fossiliferous surfaces, to constrain the life habit of Pteridinium and gain insights into the character of benthic ecosystems shortly before the beginning of the Cambrian. We present paleontological and sedimentological evidence suggesting that Pteridinium was semi-infaunal and lived gregariously in aggregated communities, preferentially adopting an orientation with the long axis perpendicular to the prevailing current direction. Using computational fluid dynamics simulations, we demonstrate that this life habit could plausibly have led to suspended food particles settling within the organism's central cavity. This supports interpretation of Pteridinium as a macroscopic suspension feeder that functioned similarly to the coeval erniettomorph Ernietta, emblematic of a broader paleoecological shift toward benthic suspension-feeding strategies over the course of the latest Ediacaran. Finally, we discuss how this new reconstruction of Pteridinium provides information concerning its potential relationships with extant animal groups and state a case for reconstructing Pteridinium as a colonial metazoan.

This work describes and illustrates Ediacaran (latest Precambrian) body and trace fossils collected in Namibia with the assistance of the Geological Survey of Namibia during 1993–1996. All of the fossils are impressions left in sandstones by the remains or activities of soft-bodied animals that have no obvious living counterparts. The challenge has been to understand the morphology of these organisms, describe their anatomy, and find places for them in the tree of life. The focus is on three erniettomorphs, Ernietta, Pteridinium, and Swartpuntia; a problematical organism named Archaeichnium that may be related to sea anemones; a new simple unmineralized sponge (Arimasia); and tubular fossils and trace fossils, all attributable to worms. We show how these fossils fit into the well-established stratigraphic context of the Nama sedimentary basin and briefly comment on their importance for the evolution of early animal life. Ediacaran fossils, obtained in stratigraphic context in 1993, 1995, and 1996, with the assistance of A. Seilacher, IGCP project 320 scientists, and the Geological Survey of Namibia, are described for the first time. Most are from the Kliphoek and Buchholzbrunn members of the Dabis Formation and the Huns and Spitskop members of the Urusis Formation, Witputs subbasin, but a significant number, including Pteridinium, are from the Kliphoek Member, Zaris Formation, and the Neiderhagen Member, Nudaus Formation, north of the Osis arch, which separates the two subbasins. We extend the stratigraphic ranges and geographic distributions of several important taxa, including Archaeichnium, Ernietta, Pteridinium, and Swartpuntia, provide reassessments of the paleobiology of these and other organisms, and describe a new sponge—possibly an unmineralized archaeocyath—Arimasia germsi n. gen. n. sp. We also describe and illustrate various ichnofossils (including the oldest known traces from the Nama Group), narrow down the first appearance of Treptichnus in the Nama succession, and reinforce the idea that there was a prolific infauna of micrometazoans during the latest Ediacaran by naming and describing previously reported microburrows found on the surfaces of gutter casts as Ariichnus vagus n. igen. n. isp.

The Ediacara biota include macroscopic, morphologically complex soft-bodied organisms that appear globally in the late Ediacaran Period (575-542 Ma). The physiology, feeding strategies, and functional morphology of the modular Ediacara organisms (rangeomorphs and erniettomorphs) remain debated but are critical for understanding their ecology and phylogeny. Their modular construction triggered numerous hypotheses concerning their likely feeding strategies, ranging from micro-to-macrophagus feeding to photoautotrophy to osmotrophy. Macrophagus feeding in rangeomorphs and erniettomorphs is inconsistent with their lack of oral openings, and photoautotrophy in rangeomorphs is contradicted by their habitats below the photic zone. Here, we combine theoretical models and empirical data to evaluate the feasibility of osmotrophy, which requires high surface area to volume (SA/V) ratios, as a primary feeding strategy of rangeomorphs and erniettomorphs. Although exclusively osmotrophic feeding in modern ecosystems is restricted to microscopic bacteria, this study suggests that (i) fractal branching of rangeomorph modules resulted in SA/V ratios comparable to those observed in modern osmotrophic bacteria, and (ii) rangeomorphs, and particularly erniettomorphs, could have achieved osmotrophic SA/V ratios similar to bacteria, provided their bodies included metabolically inert material. Thus, specific morphological adaptations observed in rangeomorphs and erniettomorphs may have represented strategies for overcoming physiological constraints that typically make osmotrophy prohibitive for macroscopic life forms. These results support the viability of osmotrophic feeding in rangeomorphs and erniettomorphs, help explain their taphonomic peculiarities, and point to the possible importance of earliest macroorganisms for cycling dissolved organic carbon that may have been present in abundance during Ediacaran times.

Ediacara-type fossils are found in a diverse array of preservational styles, implying that multiple taphonomic mechanisms might have been responsible for their preservational expression. For many Ediacara fossils, the “death mask” model has been invoked as the primary taphonomic pathway. The key to this preservational regime is the replication or sealing of sediments around the degrading organisms by microbially induced precipitation of authigenic pyrite, leading toward fossil preservation along bedding planes. Nama-style preservation, on the other hand, captures Ediacaran organisms as molds and three-dimensional casts within coarse-grained mass flow beds, and has been previously regarded as showing little or no evidence of a microbial preservational influence. To further understand these two seemingly distinct taphonomic pathways, we investigated the three-dimensionally preserved Ediacaran fossil Pteridinium simplex from mass flow deposits of the upper Kliphoek Member, Dabis Formation, Kuibis Subgroup, southern Namibia. Our analysis, using a combination of petrographic and micro-analytical methods, shows that Pteridinium simplex vanes are replicated with minor pyrite, but are most often represented by open voids that can be filled with secondary carbonate material; clay minerals are also found in association with the vanes, but their origin remains unresolved. The scarcity of pyrite and the development of voids are likely related to oxidative weathering and it is possible that microbial activities and authigenic pyrite may have contributed to the preservation of Pteridinium simplex; however, any microbes growing on P. simplex vanes within mass flow deposits were unlikely to have formed thick mats as envisioned in the death mask model. Differential weathering of replicating minerals and precipitation of secondary minerals greatly facilitate fossil collection and morphological characterization by allowing Pteridinium simplex vanes to be parted from the massive hosting sandstone.

Discovery of fossils of the Ediacara biota near the top of the Spitzkopf Member at farm Swartpunt extends the known range of these remains in Namibia more than 600 m to near the sub-Cambrian unconformity. The fossiliferous beds occur approximately 100 m above a volcanic ash dated at 543 ± 1 Ma, and thus may be the youngest Proterozoic Ediacara-type fossils reported anywhere in the world. Fossils are preserved within and on the tops of dm-thick beds of storm-deposited sandstone at two stratigraphic levels; the environment is interpreted as open marine, generally calm but with episodic disruptions by storm waves, and probably within the euphotic zone. The presence of Pteridinium carolinaense (St. Jean), which is also known from the classic sections in Ediacara and the White Sea among others, reinforces evidence from geochronology and chemostratigraphy that the Swartpunt section is terminal Neoproterozoic in age. The new genus and species Swartpuntia germsi is a large, multifoliate frond that exhibits at least three quilted petaloids. Macroscopically, Swartpuntia resembles Pteridinium and Ediacara-type fronds such as Charniodiscus traditionally interpreted as Cnidaria, whereas microscopically it exhibits segmentation that is remarkably similar to that of the putative worm Dickinsonia. Combination of diagnostic characters of these supposedly disparate groups in a single species suggests that many species of quilted Ediacaran organisms were more similar to each other than they were to any modern groups, and provides support for the concept of the “Vendobionta” as a late Neoproterozoic group of mainly multifoliate organisms with a distinctive quilted segmentation.

The youngest formations of the Neoproterozoic Windermere Supergroup in northwestern Canada (Gametrail, Blueflower, and Risky formations) record the transition from slope to shelf deposition on a prograding passive margin. Eleven facies associations are recognized, representing environments ranging from carbonate- and siliciclastic- dominated continental slope to open carbonate shelf and siliciclastic shoreface. Seven simple sequences are recognized, which can be grouped into three composite sequences. Combination of the data presented here with previous work on underlying and overlying formations indicates that the sequence-stratigraphic record is least detailed in the deepest-water facies and most detailed in shelf facies, reflecting the relative inability of high-frequency relative sea-level oscillations to affect deposition in deep-water settings. Falling-stage deposits are especially common in the upper slope region. Several major sequence boundaries (unconformities) are clustered in the interval a short distance below the Precambrian-Cambrian boundary. The most significant of these occurs high in the Blueflower Formation, not at the top of the Risky Formation as commonly inferred. This interval containing several surfaces may reflect thermal uplift related to the rifting recorded in rocks of this age in the southern Canadian Cordillera. Renewed subsidence (thermal relaxation) commenced just prior to the Neoproterozoic-Cambrian boundary, giving rise to a thick succession of shelf to nonmarine basal-Cambrian deposits. Ediacaran body fossils previously reported from the studied units occur in a range of slope to shoreface environments, including some facies that were deposited below the photic zone. The most common taxa occur across a spectrum of facies and were apparently ecological generalists.