Anatomy of the Ediacaran rangeomorph Charnia masoni.

The Ediacaran macrofossil Charnia masoni Ford is perhaps the most iconic member of the Rangeomorpha: a group of seemingly sessile, frondose organisms that dominates late Ediacaran benthic, deep-marine fossil assemblages. Despite C. masoni exhibiting broad palaeogeographical and stratigraphical ranges, there have been few morphological studies that consider the variation observed among populations of specimens derived from multiple global localities. We present an analysis of C. masoni that evaluates specimens from the UK, Canada and Russia, representing the largest morphological study of this taxon to date. We describe substantial morphological variation within C. masoni and present a new morphological model for this species that has significant implications both for interpretation of rangeomorph architecture, and potentially for existing taxonomic schemes. Previous reconstructions of Charnia include assumptions regarding the presence of structures seen in other rangeomorphs (e.g. an internal stalk) and of homogeneity in higher order branch morphology; observations that are not borne out by our investigations. We describe variation in the morphology of third and fourth order branches, as well as variation in gross structure near the base of the frond. The diagnosis of Charnia masoni is emended to take account of these new features. These findings highlight the need for large-scale analyses of rangeomorph morphology in order to better understand the biology of this long-enigmatic group.NER

Anatomy of the Ediacaran rangeomorph Charnia masoni

The Ediacaran macrofossil Charnia masoni Ford is perhaps the most iconic member of the Rangeomorpha: a group of seemingly sessile, frondose organisms that dominates late Ediacaran benthic, deep‐marine fossil assemblages. Despite C. masoni exhibiting broad palaeogeographical and stratigraphical ranges, there have been few morphological studies that consider the variation observed among populations of specimens derived from multiple global localities. We present an analysis of C. masoni that evaluates specimens from the UK, Canada and Russia, representing the largest morphological study of this taxon to date. We describe substantial morphological variation within C. masoni and present a new morphological model for this species that has significant implications both for interpretation of rangeomorph architecture, and potentially for existing taxonomic schemes. Previous reconstructions of Charnia include assumptions regarding the presence of structures seen in other rangeomorphs (e.g. an internal stalk) and of homogeneity in higher order branch morphology; observations that are not borne out by our investigations. We describe variation in the morphology of third and fourth order branches, as well as variation in gross structure near the base of the frond. The diagnosis of Charnia masoni is emended to take account of these new features. These findings highlight the need for large‐scale analyses of rangeomorph morphology in order to better understand the biology of this long‐enigmatic group

The developmental biology of <i>Charnia</i> and the eumetazoan affinity of the Ediacaran rangeomorphs.

Molecular timescales estimate that early animal lineages diverged tens of millions of years before their earliest unequivocal fossil evidence. The Ediacaran macrobiota (~574 to 538 million years ago) are largely eschewed from this debate, primarily due to their extreme phylogenetic uncertainty, but remain germane. We characterize the development of Charnia masoni and establish the affinity of rangeomorphs, among the oldest and most enigmatic components of the Ediacaran macrobiota. We provide the first direct evidence for the internal interconnected nature of rangeomorphs and show that Charnia was constructed of repeated branches that derived successively from pre-existing branches. We find homology and rationalize morphogenesis between disparate rangeomorph taxa, before producing a phylogenetic analysis, resolving Charnia as a stem-eumetazoan and expanding the anatomical disparity of that group to include a long-extinct bodyplan. These data bring competing records of early animal evolution into closer agreement, reformulating our understanding of the evolutionary emergence of animal bodyplans

The tempo of Ediacaran evolution

The rise of complex macroscopic life occurred during the Ediacaran Period, an interval that witnessed large-scale disturbances to biogeochemical systems. The current Ediacaran chronostratigraphic framework is of insufficient resolution to provide robust global correlation schemes or test hypotheses for the role of biogeochemical cycling in the evolution of complex life. Here, we present new radio-isotopic dates from Ediacaran strata that directly constrain key fossil assemblages and large-magnitude carbon cycle perturbations. These new dates and integrated global correlations demonstrate that late Ediacaran strata of South China are time transgressive and that the 575- to 550-Ma interval is marked by two large negative carbon isotope excursions: the Shuram and a younger one that ended ca. 550 Ma ago. These data calibrate the tempo of Ediacaran evolution characterized by intervals of tens of millions of years of increasing ecosystem complexity, interrupted by biological turnovers that coincide with large perturbations to the carbon cycle

A reassessment of the problematic Ediacaran genus Orbisiana Sokolov 1976

The genus Orbisiana was established in 1976 by B.S. Sokolov based on a collection of seven specimens within core material from a borehole drilled through the Ediacaran-age Gavrilov Yam Formation of the Moscow Basin, Russia. Here we reassess the original material for the type species Orbisiana simplex Sokolov 1976, which was long considered to be lost; fix the holotype of the type species; and revise the original diagnosis of the genus and species. Pyritisation of the fossils, which are preserved in finely laminated shales, allows three-dimensional morphological characterisation of this taxon using X-ray microtomography (μCT). Morphological and taphonomic analyses of the type material and additional three-dimensionally preserved specimens from the Verkhovka Formation (Vendian of the White Sea area) suggest that Orbisiana simplex consisted of submillimetric to millimetric globular chambers arranged in compact, grape-like clusters, or forming sinuous to linear aggregates. Occasionally, aggregates can bifurcate, with no appreciable change in chamber dimensions or shape. The phylogenetic affinity of Orbisiana remains uncertain, but its chambered construction, putative agglutinated structure of the chamber walls, and compact, occasionally branching chamber arrangement are shared with agglutinated tests of the Ediacaran genus Palaeopascichnus. Our reassessment and systematic study of the genus Orbisiana sheds new light on one of the least studied members of the late Ediacaran macroscopic biota

Arumberiamorph structure in modern microbial mats: implications for Ediacaran palaeobiology

In the course of studying modern halotolerant microbial mats in salterns near the village of Kervalet, western France, we observed fanning-out and curved series of macroscopic ridges on the surface of a newly formed biofilm. The structure resembles the late Ediacaran fossil Arumberia which is globally distributed in Australia, Avalonia, Baltica, Siberia and India, always confined to intertidal and delta-plain settings subject to periodic desiccation or fluctuating salinity. Although the origin of the structure observed in modern microbial mats remains enigmatic, wrinkled and rugose variants of microbial biofilms in general exhibit increased levels of resistance to several environmental stresses. By analogy, the fossil Arumberia could be interpreted as a microbial mat morphotype (the “Arumberia” morph) developed in response to environmental perturbations in terminal Ediacaran shallow marine basins. If environmental conditions are likely to be responsible for the formation of Arumberia, it is not that a specific biological community has survived since the Ediacaran – it is that the biological response of microbial communities that manifested itself quite commonly in certain terminal Ediacaran and early Cambrian environments can still be found (seemingly in much more restricted settings) today

Three-dimensional reconstruction, taphonomic and petrological data suggest the oldest record of bioturbation is a body fossil coquina

Fossil material assigned to Nenoxites from the late Ediacaran Khatyspyt Formation of Arctic Siberia (550–544 Ma) has been presented as evidence for bioturbation prior to the basal Cambrian boundary. However, that ichnological interpretation has been challenged, and descriptions of similar material from other global localities support a body fossil origin. Here we combine X-ray computed tomography (μCT), scanning electron microscopy (SEM), and petrographic methods to evaluate the body- or tracefossil nature of Nenoxites from the Khatyspyt Formation. The fossilised structures comprise densely-packed chains of three-dimensionally preserved silicic, bowl-shaped elements surrounded by distinct sedimentary halos, within a dolomitized matrix. Individual bowl-shaped elements can exhibit diffuse mineralogical boundaries and bridging connections between elements, both considered here to result from silicification and dolomitization during diagenesis. This new morphological and petrological evidence, in conjunction with recent studies of the late Ediacaran tubular taxa Ordinilunulatus and Shaanxilithes from China, leads us to conclude that the Khatyspyt specimens most likely reflect a coquina deposit of Shaanxilithes-like body fossils. Our data support the possibility of Shaanxilithes-like organisms representing total group eumetazoans.NERC (NE/L011409/1 and NE/P013678/1 Leverhulme Trust Research Fellowship (RF-2022-167). Early career fellowship from St. Edmund Hall, Oxford. Government Contract FWZZ-2022-0002 (Fundamental Scientific Research Programme of the Russian Federation). Russian Science Foundation (grant 20-67-46028

Understanding the geobiology of the terminal Ediacaran Khatyspyt Lagerstätte (Arctic Siberia, Russia)

The Khatyspyt Lagerstätte (~544 Ma, Russia) provides a valuable window into late Ediacaran Avalon-type ecosystems with rangeomorphs, arboreomorphs, and mega-algae. Here, we tackle the geobiology of this Lagerstätte by the combined analysis of paleontological features, sedimentary facies, and lipid biomarkers. The Khatyspyt Formation was deposited in carbonate ramp environments. Organic matter (0.12–2.22 wt.% TOC) displays characteristic Ediacaran biomarker features (e.g., eukaryotic steranes dominated by the C29 stigmastane). Some samples contain a putative 2-methylgammacerane that was likely sourced by ciliates and/or bacteria. 24-isopropylcholestane and 26-methylstigmastane are consistently scarce (≤0.4% and ≤0.2% of ∑C27-30 regular steranes, respectively). Thus, Avalon-type organisms occupied different niches than organisms capable of directly synthesizing C30 sterane precursors among their major lipids. Relative abundances of eukaryotic steranes and bacterial hopanes (sterane/hopane ratios = 0.07–0.30) demonstrate oligotrophic and bacterially dominated marine environments, similar to findings from other successions with Ediacara-type fossils. Ediacara-type fossils occur in facies characterized by microbial mats and biomarkers indicative for a stratified marine environment with normal–moderate salinities (moderate–high gammacerane index of 2.3–5.7; low C35 homohopane index of 0.1–0.2). Mega-algae, in contrast, are abundant in facies that almost entirely consist of allochthonous event layers. Biomarkers in these samples indicate a non-stratified marine environment and normal salinities (low gammacerane index of 0.6–0.8; low C35 homohopane index of 0.1). Vertical burrowers occur in similar facies but with biomarker evidence for stratification in the water column or around the seafloor (high gammacerane index of 5.6). Thus, the distribution of macro-organisms and burrowers was controlled by various, dynamically changing environmental factors. It appears likely that dynamic settings like the Khatyspyt Lagerstätte provided metabolic challenges for sustenance and growth which primed eukaryotic organisms to cope with changing environmental habitats, allowing for a later diversification and expansion of complex macroscopic life in the marine realm

Data from: Anatomy of the Ediacaran rangeomorph Charnia masoni

The Ediacaran macrofossil Charnia masoni Ford is perhaps the most iconic member of the Rangeomorpha: a group of seemingly sessile, frondose organisms that dominates late Ediacaran benthic, deep‐marine fossil assemblages. Despite C. masoni exhibiting broad palaeogeographical and stratigraphical ranges, there have been few morphological studies that consider the variation observed among populations of specimens derived from multiple global localities. We present an analysis of C. masoni that evaluates specimens from the UK, Canada and Russia, representing the largest morphological study of this taxon to date. We describe substantial morphological variation within C. masoni and present a new morphological model for this species that has significant implications both for interpretation of rangeomorph architecture, and potentially for existing taxonomic schemes. Previous reconstructions of Charnia include assumptions regarding the presence of structures seen in other rangeomorphs (e.g. an internal stalk) and of homogeneity in higher order branch morphology; observations that are not borne out by our investigations. We describe variation in the morphology of third and fourth order branches, as well as variation in gross structure near the base of the frond. The diagnosis of Charnia masoni is emended to take account of these new features. These findings highlight the need for large‐scale analyses of rangeomorph morphology in order to better understand the biology of this long‐enigmatic group.,Charnia masoni Holotype RTISupplementary Figure 1: Reflectance transformation image of the Charnia masoni holotype (LEIUG 2328).