Jack J. Matthews

Haootia quadriformis n. gen., n. sp., interpreted as a muscular cnidarian impression from the Late Ediacaran period (approx. 560 Ma)

Muscle tissue is a fundamentally eumetazoan attribute. The oldest evidence for fossilized muscular tissue before the Early Cambrian has hitherto remained moot, being reliant upon indirect evidence in the form of Late Ediacaran ichnofossils. We here report a candidate muscle-bearing organism, Haootia quadriformis n. gen., n. sp., from approximately 560 Ma strata in Newfoundland, Canada. This taxon exhibits sediment moulds of twisted, superimposed fibrous bundles arranged quadrilaterally, extending into four prominent bifurcating corner branches. Haootia is distinct from all previously published contemporaneous Ediacaran macrofossils in its symmetrically fibrous, rather than frondose, architecture. Its bundled fibres, morphology, and taphonomy compare well with the muscle fibres of fossil and extant Cnidaria, particularly the benthic Staurozoa. Haootia quadriformis thus potentially provides the earliest body fossil evidence for both metazoan musculature, and for Eumetazoa, in the geological record.

A new assemblage of juvenile Ediacaran fronds from the Drook Formation, Newfoundland

A new assemblage of frondose and filamentous Ediacaran macrofossils is reported from the upper Drook Formation of Pigeon Cove, Newfoundland. The frondose forms, all less than 3 cm in length, are considered to represent the juvenile growth stages of Ediacaran organisms including Charnia spp. and Trepassia spp. This is the first report of an assemblage wholly dominated by such small juvenile rangeomorph forms, and provides insights into the ontogeny and ecology of these earliest members of the Ediacara biota. The fronds occur alongside filamentous forms with similarities to microbial taxa, and both morphotypes are considered to postdate an assemblage of large ivesheadiomorphs on the same bedding plane. If so, the assemblage represents one of the oldest documented examples of secondary community succession. The new Pigeon Cove fossils also extend the stratigraphic ranges of several key frondose taxa (Charnia masoni, Charniodiscus spp.) back into some of the oldest known macrofossil-bearing strata. These revised ranges lend support to the suggestion that the previously observed low diversity within the Drook Formation may represent a combination of taphonomic and sampling artefacts. Furthermore, this assemblage implies that the diversification of architectural morphotypes within the Ediacara biota took place earlier than hitherto suspected. Supplementary material: A document containing figures of additional juvenile rangeomorphs and filamentous specimens, a table of specimen dimensions, and a complete digitized map of the Pigeon Cove bedding plane, is available at www.geolsoc.org.uk/SUP18529.

Uncovering framboidal pyrite biogenicity using nano-scale CNorg mapping

Framboidal pyrite has been used as a paleo-redox proxy and a biomarker in ancient sediments, but the interpretation of pyrite framboids can be controversial, especially where later overgrowths have obscured primary textures. Here we show how nano-scale chemical mapping of organic carbon and nitrogen (CN org ) can detect relict framboids within Precambrian pyrite grains and determine their formation mechanism. Pyrite grains associated with an Ediacaran fossil Lagerstatte from Newfoundland (ca. 560 Ma) hold significance for our understanding of taphonomy and redox history of the earliest macrofossil assemblages. They show distinct chemical zoning with respect to CN org . Relict framboids are revealed as spheroidal zones within larger pyrite grains, whereby pure pyrite microcrystals are enclosed by a mesh-like matrix of pyrite possessing elevated CN org , replicating observations from framboids growing within modern biofilms. Subsequent pyrite overgrowths also incorporated CN org from biofilms, with concentric CN org zoning showing that the availability of CN org progressively decreased during later pyrite growth. Multiple framboids are commonly cemented together by these overgrowths to form larger grains, with relict framboids only detectable in CN org maps. In situ sulfur isotope data (δ 34 S = ∼−24‰ to −15‰) show that the source of sulfur for the pyrite was also biologically mediated, most likely via a sulfate-reducing microbial metabolism within the biofilms. Relict framboids have significantly smaller diameters than the pyrite grains that enclose them, suggesting that the use of framboid diameters to infer water column paleo-redox conditions should be approached with caution. This work shows that pyrite framboids have formed within organic biofilms for at least 560 m.y., and provides a novel methodology that could readily be extended to search for such biomarkers in older rocks and potentially on other planets.

Reconstructing the reproductive mode of an Ediacaran macro-organism

Enigmatic macrofossils of late Ediacaran age (580–541 million years ago) provide the oldest known record of diverse complex organisms on Earth, lying between the microbially dominated ecosystems of the Proterozoic and the Cambrian emergence of the modern biosphere. Among the oldest and most enigmatic of these macrofossils are the Rangeomorpha, a group characterized by modular, self-similar branching and a sessile benthic habit. Localized occurrences of large in situ fossilized rangeomorph populations allow fundamental aspects of their biology to be resolved using spatial point process techniques. Here we use such techniques to identify recurrent clustering patterns in the rangeomorph Fractofusus, revealing a complex life history of multigenerational, stolon-like asexual reproduction, interspersed with dispersal by waterborne propagules. Ecologically, such a habit would have allowed both for the rapid colonization of a localized area and for transport to new, previously uncolonized areas. The capacity of Fractofusus to derive adult morphology by two distinct reproductive modes documents the sophistication of its underlying developmental biology.

Confirming the metazoan character of a 565 Ma trace-fossil assemblage from Mistaken Point, Newfoundland

ABSTRACT Surface locomotory trace fossils from the Mistaken Point Formation of Newfoundland, dated at ∼ 565 Ma, suggest that organisms capable of controlled locomotion and possessing muscular tissue may have existed among Avalonian Ediacaran macrofossil assemblages. Here we describe the Mistaken Point trace-fossil assemblage in full, discuss its stratigraphic context within the Mistaken Point Formation, and explore the competing hypotheses for the formation of the traces. We find that the trace fossils, preserved within a turbidite succession in a deep-marine depositional environment, are not attributable to abiogenic structures, to Ediacaran tubular or filamentous body fossils, to rangeomorph stems, or to a host of late Ediacaran and early Phanerozoic ichnofossils. Specimens within the assemblage show some similarities to the ichnogenera Helminthoidichnites and Archaeonassa, but discrepancies in certain aspects of their structure mean that we do not formally attribute them to these ichnotaxa at this time. The Mistaken Point ichnofossils possess morphological characteristics indicative of formation by an organism with a round base. Comparison with traces formed by modern organisms of such character appears to rule out formation by protistan, echinoderm, or annelid styles of movement, but is consistent with organisms moving via muscular controlled locomotion in a similar way to some modern mollusks and actinian cnidarians. We suggest therefore that the Mistaken Point trace-fossil assemblage reveals the presence of muscular metazoans in late Ediacaran deep-marine ecosystems. Such organisms cannot yet be attributed to specific phyla, but their inferred locomotory mechanisms share closest similarities with those utilized by extant actinians.

Post-fossilization processes and their implications for understanding Ediacaran macrofossil assemblages

Abstract Fossil assemblages from Newfoundlands Avalon Peninsula preserve diverse examples of the enigmatic Ediacaran macrobiota, offering some of the earliest evidence for large and complex multicellular life. These fossils are exposed on extensive coastal bedding planes in extraordinary abundances, permitting palaeoecological studies based on census data from spatially extensive palaeocommunities. Such studies have been used to constrain the reproductive strategy and phylogenetic placement of Ediacaran organisms. Geological mapping and stratigraphic correlation in the Mistaken Point Ecological Reserve reveal that some fossil-bearing surfaces can be tracked over distances of several kilometres. These laterally extensive surfaces reveal that the modern processes by which the sediment overlying a fossil surface is removed may impose important controls on the observed composition of fossil assemblages. Weathering and erosion – along with factors associated with tectonics, metamorphism and discovery – are here grouped as ‘post-fossilization processes’ and introduce biases that are often not explicitly accounted for in palaeoecological studies. Specifically, post-fossilization processes may differentially influence the preservational fidelity of individual specimens on a given surface and generate features that could be mistaken for original morphological characters. We therefore recommend that post-fossilization processes must be considered when undertaking palaeoecological studies in Ediacaran successions in Newfoundland and, potentially, elsewhere.

The arrangement of possible muscle fibres in the Ediacaran taxon Haootia quadriformis

Haootia quadriformis from Newfoundland, Canada, is one of the most unusual impressions of a soft-bodied macro-organism yet described from the late Ediacaran Period. Interpreted as a metazoan of cnidarian grade [[1][1]], the body impression of H. quadriformis possesses features interpreted as fibrous