The utility of height for the Ediacaran organisms of Mistaken Point.

Ediacaran fossil communities consist of the oldest macroscopic eukaryotic organisms. Increased size (height) is hypothesized to be driven by competition for water column resources, leading to vertical/epifaunal tiering and morphological innovations such as stems. Using spatial analyses, we find no correlation between tiering and resource competition, and that stemmed organisms are not tiered. Instead, we find that height is correlated with greater offspring dispersal, demonstrating the importance of colonization potential over resource competition.Gibbs Travelling Fellowship from Newnham College, University of Cambridge Henslow Research Fellowship, Cambridge Philosophical Society

Of time and taphonomy: preservation in the Ediacaran

The late Neoproterozoic witnessed a revolution in the history of life: the transition from a microbial world to the one we know today. The enigmatic organisms of the Ediacaran hold the key to understanding the early evolution of metazoans and their ecology, and thus the basis of Phanerozoic life. Crucial to interpreting the information they divulge is a thorough understanding of their taphonomy: of what is preserved andhow it is preserved, and also of what is not preserved. Fortunately, this Period is also recognized for its abundance of soft-tissue preservation, which is viewed through a wide variety of taphonomic windows. Some of these, such as pyritization and carbonaceous compression, are also present throughout the Phanerozoic, but the abundance and variety of moldic preservation of body fossils in siliclastic settings is unique to the Ediacaran. In rare cases, one organism is preserved in several preservational styles which, in conjunction with our increased understanding of the taphonomic processes involved in each style, allow us to more confidently interpret aspects of the biology and ecology of the organisms preserved. Several groundbreaking advances in this field have been made since the 1990s, and have paved the way for increasingly thorough analyses and elegant interpretations

The Palaeobiology of Avalonian (Ediacaran) Rangeomorphs

The Earth has supported life for most of its 4.5 billion year history, but the first macroscopic organisms only appeared some 600 million years ago, in the Ediacaran. Their world was fundamentally different from the one we know today, and many aspects of their biology and ecology remain a mystery. The late Ediacaran fossil assemblages of Avalonia represent some of the oldest evidence for complex macroscopic life, and are dominated by rangeomorphs, a group characterised by their self-similar branching architecture. In this thesis, I investigate several aspects of the preservation, classification and ecology of these enigmatic deep marine organisms. The biotas of Charnwood Forest host several taxa which are new to science. Five of these are described here, and include two new genera, Orthiokaterna fordi gen. et sp. nov., and Undosyrus nemoralis gen. et sp. nov., and three new species: Primocandelabrum anatonos sp. nov., P. boytoni sp. nov., and P. katatonos sp. nov.. The Primocandelabrum species in particular encompass a great deal of variation in both branching characters and overall morphology. By using a novel multivariate statistical approach to analyse multiple characters in tandem, individual taxa can be discriminated from one another. Much of the observed variation is interpreted as intra-specific. This level of variation within a single taxon has not previously been recognised in rangeomorphs, and is likely attributable to (eco?)phenotypic rather than ontogenetic variability. Orthiokaterna displays eccentric branches, interpreted as a growth response to mechanical damage, reflecting a greater degree of growth plasticity than that recognised in other rangeomorphs, while Undosyrus had an external sheath, interpreted as modified rangeomorph elements serving a protective role. Even without knowing the phylogenetic relationships of rangeomorphs, it is possible to resolve key aspects of their palaeoecology. The response(s) of communities in Charnwood Forest and Newfoundland to both ambient disturbance and to more substantial events is investigated by combining detailed petrographic analysis of the host sediments with multivariate statistical techniques. I demonstrate that higher taxonomic diversity is correlated with low–intermediate physical disturbance; that upright taxa (e.g. Charnia) dominate surfaces which experienced small-scale, sub-lethal sedimentation events and comparably high background sediment input; and that flat-lying forms (e.g. Fractofusus) preferentially occur on surfaces with low sediment input. The population demographics of several taxa also show evidence of multimodality: in some (including Charnia and Primocandelabrum), bimodality was induced by culling of part of an incumbent population by a substantial disturbance event, followed by re-colonisation; in others (e.g. Fractofusus), overlapping cohorts reflect non-continuous or pulsed reproduction. Disturbance (ambient and discrete events) demonstrably influenced community succession, with early-colonising taxa dominating horizons with low overall levels of disturbance, and those able to survive disturbance events dominating recovery populations and horizons with higher levels of disturbance. Based on their inferred life history traits and their environmental preferences, I propose a model of ecological succession for rangeomorph communities

Of time and taphonomy: preservation in the Ediacaran

The late Neoproterozoic witnessed a revolution in the history of life: the transition from a microbial world to the one known today. The enigmatic organisms of the Ediacaran hold the key to understanding the early evolution of metazoans and their ecology, and thus the basis of Phanerozoic life. Crucial to interpreting the information they divulge is a thorough understanding of their taphonomy: what is preserved, how it is preserved, and also what is not preserved. Fortunately, this Period is also recognized for its abundance of soft-tissue preservation, which is viewed through a wide variety of taphonomic windows. Some of these, such as pyritization and carbonaceous compression, are also present throughout the Phanerozoic, but the abundance and variety of moldic preservation of body fossils in siliciclastic settings is unique to the Ediacaran. In rare cases, one organism is preserved in several preservational styles which, in conjunction with an increased understanding of the taphonomic processes involved in each style, allow confident interpretations of aspects of the biology and ecology of the organisms preserved. Several groundbreaking advances in this field have been made since the 1990s, and have paved the way for increasingly thorough analyses and elegant interpretations

Modularity and overcompensatory growth in Ediacaran rangeomorphs demonstrate early adaptations for coping with environmental pressures

The first known diverse, complex, macroscopic benthic marine ecosystems (late Ediacaran, ca. 571-541 Ma) were dominated by the Rangeomorpha, an enigmatic group of extinct frondose eukaryotes that are candidate early metazoans[1,2]. The group is characterised by a self-similar branching architecture that was likely optimised for exchange, but nearly every other aspect of their biology is contentious[2-4]. We report locally-enhanced, aberrant growth ("eccentric branching") in a stalked, multifoliate rangeomorph - Hylaecullulus fordi n. gen., n. sp. - from Charnwood Forest (UK), confirming the presence of true biological modularity within the group. Random branches achieve unusually large proportions and mimic the architecture of their parent branch, rather than that of their neighbours (the norm). Their locations indicate exceptional growth at existing loci, rather than insertion at new sites. Analogous over- compensatory branching in extant modular organisms requires the capacity to orchestrate growth at specific sites, and occurs most frequently in response to damage or environmental stress, allowing regeneration towards optimum morphology[e.g. 5-7]. Its presence in rangeomorphs indicates a hitherto unappreciated level of control to their growth plan, a previously unrecognised form of morphological plasticity within the group, and an ability to actively respond to external physical stimuli. The trait would have afforded rangeomorphs resilience to fouling and abrasion, partially accounting for their wide environmental tolerance, and may have pre-adapted them to withstand predation, weakening this argument for their extinction. Our findings highlight that multiple, phylogenetically disparate, clades first achieved large size through modularity

Orientations of Mistaken Point Fronds Indicate Morphology Impacted Ability to Survive Turbulence

The Ediacaran fossils of the Mistaken Point E surface have provided crucial insight into early animal communities, including how they reproduced, the importance of Ediacaran height and what the most important factors were to their community dynamics. Here, we use this iconic community to investigate how morphological variation between eight taxa affected their ability to withstand different flow conditions. For each of Beothukis, Bradgatia, Charniodiscus procerus, Charniodiscus spinosus, Plumeropriscum, Primocandelabrum, Thectardis and Fractofusus we measured the orientation and length of their stems (if present) and their fronds. We statistically tested each taxon’s stem and frond orientation distributions to see whether they displayed a uniform or multimodal distribution. Where multimodal distributions were identified, the stem/frond length of each cohort was tested to identify if there were differences in size between different orientation groups. We find that Bradgatia and Thectardis show a bimodal felling direction, and infer that they were felled by the turbulent head of the felling flow. In contrast, the frondose rangeomorphs including Beothukis, Plumeropriscum, Primocandelabrum, and the arboreomorphs were felled in a single direction, indicating that they were upright in the water column, and were likely felled by the laminar tail of the felling flow. These differences in directionality suggests that an elongate habit, and particularly possession of a stem, lent greater resilience to frondose taxa against turbulent flows, suggesting that such taxa would have had improved survivability in conditions with higher background turbulence than taxa like Bradgatia and Thectardis, that lacked a stem and had a higher centre of mass, which may have fared better in quieter water conditions.</jats:p

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

Integrated records of environmental change and evolution challenge the Cambrian Explosion.

The 'Cambrian Explosion' describes the rapid increase in animal diversity and abundance, as manifest in the fossil record, between ~540 and 520 million years ago (Ma). This event, however, is nested within a far more ancient record of macrofossils extending at least into the late Ediacaran at ~571 Ma. The evolutionary events documented during the Ediacaran-Cambrian interval coincide with geochemical evidence for the modernisation of Earth's biogeochemical cycles. Holistic integration of fossil and geochemical records leads us to challenge the notion that the Ediacaran and Cambrian worlds were markedly distinct, and places biotic and environmental change within a longer-term narrative. We propose that the evolution of metazoans may have been facilitated by a series of dynamic and global changes in redox conditions and nutrient supply, which, potentially together with biotic feedbacks, enabled turnover events that sustained multiple phases of radiation. We argue that early metazoan diversification should be recast as a series of successive, transitional radiations that extended from the late Ediacaran and continued through the early Palaeozoic. We conclude that while the Cambrian Explosion represents a radiation of crown-group bilaterians, it was simply one phase amongst several metazoan radiations, some older and some younger

The Ediacaran fossils of Charnwood Forest: shining new light on a major biological revolution

Charnwood Forest (UK) hosts some of the oldest and best-preserved macrofossils known from the Ediacaran. It is the counterpoint to the more widely studied fossil sites of south-eastern Newfoundland (Canada), which include the recently-designated UNESCO World Heritage Site of Mistaken Point. Discoveries made in Charnwood Forest since 2008 have the potential to revolutionise our understanding of the evolution of complex macroscopic life and the subsequent development of ‘modern’ (i.e. Phanerozoic) ecosystems. The sites in Charnwood include the holotypes for several iconic Ediacaran taxa, and potentially both the oldest and youngest representatives of the deep-water Avalon Assemblage. These communities provide a unique opportunity to test models of community ecology, biological endemism and environmental sensitivity and adaptability in the Ediacaran. Here, we review the geology of Charnwood Forest and the palaeobiology of its biotas, and we summarise recent scientific advances in the context of our developing understanding of early macroscopic life. We review the application of Reflectance Transformation Imaging to these ancient communities, and signpost exciting new directions for research in Charnwood Forest, almost 170 years after the fossils were first brought to light