Role of low intensity environmental disturbance in structuring the earliest (Ediacaran) macrobenthic tiered communities

Rangeomorphs were important components of Ediacaran macrobenthic ecosystems, yet their biology and ecology remain poorly constrained. They formed high-density, tiered communities that were subjected to intermittent burial events, the largest of which killed entire communities. Abundant thin event beds in the Ediacaran succession of Charnwood Forest indicate the additional, frequent impact of minor obrution events. The type surface of Charnia masoni is immediately underlain by one such lamina (a tuff) and preserves a distinctly bimodal population. It is dominated by Charnia fronds that are of smaller or comparable length to the holotype (19.4. cm), but also includes notably larger specimens (>. 45. cm) that would traditionally have been assigned to Charnia grandis. Multiple morphological- and morphometric parameters (length, width, spacing of primary branches) demonstrate that these are indistinguishable from the holotype of C. masoni, affirming the synonymy of the two taxa. Nevertheless, these outsized individuals are distinguished by their proportionally fewer primary branches per unit length. Taphonomic evidence indicates that they were survivors of an incumbent population, the rest of which was culled by a minor ashfall. We suggest that this temporary reduction in competition from neighbours allowed the survivors to grow larger and thereby gain access to a greater proportion of the water column. As the community recovered, their large size would have continued to provide them with an advantage, divorcing them from the density-dependent competition seen in the new understory. The interlude between cohorts implies that new recruits were substrate-sensitive, presumably awaiting re-establishment of the biomat. Sub-lethal disturbance events thus played a significant role in structuring Ediacaran communities, and help explain the observed bed-by-bed variability. Taken as a whole, the growth trajectory of C. masoni resembles that of extant organisms with indeterminate growth programmes and no genetically-controlled upper size limit

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

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

U-Pb geochronology and global context of the Charnian Supergroup, UK: constraints on the age of key Ediacaran fossil assemblages

U-Pb (zircon) ages for key stratigraphic volcanic horizons within the ∼3200-m-thick Ediacaran-age Charnian Supergroup provide an improved age model for the included Avalonian assemblage macrofossils and, hence, temporal constraints essential for intercomparisons of the Charnian fossils with other Ediacaran fossil assemblages globally. The Ives Head Formation (Blackbrook Group), the oldest exposed part of the volcaniclastic Charnian Supergroup of the late Neoproterozoic Avalonian volcanic arc system of southern Britain, contains a bedding plane with an impoverished assemblage of ivesheadiomorphs that is constrained to between ca. 611 Ma and 569.1 ± 0.9 Ma (total uncertainty). Higher-diversity biotas, including the holotypes of Charnia, Charniodiscus, and Bradgatia, occupy the upper part of the volcaniclastic succession (Maplewell Group) and are dated at 561.9 ± 0.9 Ma (total uncertainty) and younger by zircons interpreted as coeval with eruption and deposition of the Park Breccia, Bradgate Formation. An ashy volcanic-pebble conglomerate in the Hanging Rocks Formation at the very top of the supergroup yielded two U-Pb zircon populations: an older detrital one at ca. 604 Ma, and a younger population at ca. 557 Ma, which is interpreted as the approximate depositional age. The temporal association of the fossiliferous Charnian Supergroup with comparable fossiliferous deep-water successions in Newfoundland, and the probable temporal overlap of the youngest Charnwood macrofossils with those from different paleoenvironmental settings, such as the Ediacaran White Sea macrofossils, indicate a primary role for ecological sensitivity in determining the composition of these late Neoproterozoic communities

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

Polymorphic organization in a planktonic graptoloid (Hemichordata: Pterobranchia) colony of Late Ordovician age

Graptolites are common fossils in Early Palaeozoic strata, but little is known of their soft-part anatomy. However, we report a long-overlooked specimen of Dicranograptus aff. ramosus from Late Ordovician strata of southern Scotland that preserves a strongly polymorphic, recalcitrant, organic-walled network hitherto unseen in graptoloid graptolites. This network displays three morphologies: proximally, a strap-like pattern, likely of flattened tubes; these transform distally into isolated, hourglass-shaped structures; then, yet more distally, revert to a (simpler) strap-like pattern. The network most likely represents a stolon-like system, hitherto unknown in graptoloids, that connected individual zooids. Its alternative interpretation, as colonial xenobionts that infested a graptoloid colony and mimicked its architecture, is considered less likely on taphonomic and palaeobiological grounds. Such polymorphism is not known in non-graptolite pterobranchs, which are less diverse and morphologically more conservative: a division of labour between graptoloid zooids for such functions as feeding, breeding and rhabdosome construction may have been the key to their remarkable evolutionary success

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 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

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

A combined geomorphological and geophysical approach to characterising relict landslide hazard on the Jurassic Escarpments of Great Britain

The Jurassic Escarpment in the North York Moors in Northern Britain has a high density of deep-seated relict landslides but their regional hazard is poorly understood due to a lack of detailed case studies. Investigation of a typical relict landslide at Great Fryup Dale suggests that the crop of the Whitby Mudstone Formation is highly susceptible to landslide hazards. The mudstone lithologies along the Escarpment form large multiple rotational failures which break down at an accelerated rate during wetter climates and degrade into extensive frontal mudflows. Geomorphological mapping, high resolution LiDAR imagery, boreholes, and geophysical ERT surveys are deployed in a combined approach to delimit internal architecture of the landslide. Cross-sections developed from these data indicate that the main movement displaced a bedrock volume of c. 1 × 107 m3 with a maximum depth of rupture of c. 50 m. The mode of failure is strongly controlled by lithology, bedding, joint pattern, and rate of lateral unloading. Dating of buried peats using the AMS method suggests that the 10 m thick frontal mudflow complex was last active in the Late Holocene, after c. 2270 ± 30 calendar years BP. Geomorphic mapping and dating work indicates that the landslide is dormant, but slope stability modelling suggests that the slope is less stable than previously assumed; implying that this and other similar landslides in Britain may become more susceptible to reactivation or extension during future wetter climatic phases. This study shows the value of a multi-technique approach for landslide hazard assessment and to enhance national landslide inventories