No abstract

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.

The Conception and St. John’s Groups of southeastern Newfoundland contain some of the oldest known fossils of the Ediacaran macrobiota. The Mistaken Point Ecological Reserve UNESCO World Heritage Site is an internationally recognized locality for such fossils and hosts early evidence for both total group metazoan body fossils and metazoan-style locomotion. The Mistaken Point Ecological Reserve sedimentary succession includes ∼1500 m of fossil-bearing strata containing numerous dateable volcanogenic horizons, and therefore offers a crucial window into the rise and diversification of early animals. Here we present six stratigraphically coherent radioisotopic ages derived from zircons from volcanic tuffites of the Conception and St. John’s Groups at Mistaken Point Ecological Reserve. The oldest architecturally complex macrofossils, from the upper Drook Formation, have an age of 574.17 ± 0.66 Ma (including tracer calibration and decay constant uncertainties). The youngest rangeomorph fossils from Mistaken Point Ecological Reserve, in the Fermeuse Formation, have a maximum age of 564.13 ± 0.65 Ma. Fossils of the famous “E” Surface are confirmed to be 565.00 ± 0.64 Ma, while exceptionally preserved specimens on the “Brasier” Surface in the Briscal Formation are dated at 567.63 ± 0.66 Ma. We use our new ages to construct an age-depth model for the sedimentary succession, constrain sedimentary accumulation rates, and convert stratigraphic fossil ranges into the time domain to facilitate integration with time-calibrated data from other successions. Combining this age model with compiled stratigraphic ranges for all named macrofossils within the Mistaken Point Ecological Reserve succession, spanning 76 discrete fossil-bearing horizons, enables recognition and interrogation of potential evolutionary signals. Peak taxonomic diversity is recognized within the Mistaken Point and Trepassey Formations, and uniterminal rangeomorphs with undisplayed branching architecture appear several million years before multiterminal, displayed forms. Together, our combined stratigraphic, paleontological, and geochronological approach offers a holistic, time-calibrated record of evolution during the mid−late Ediacaran Period and a framework within which to consider other geochemical, environmental, and evolutionary data sets.

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 Lagerstätte 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 (d 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.