Ediacaran biota

The first Ediacaran fossils discovered were the disc-shaped Aspidella terranovica in 1868. Their discoverer, Scottish geologist Alexander Murray, found them useful aids for correlating the age of rocks around Newfoundland. In 1933, Georg Gürich discovered specimens in Namibia but assigned them to the Cambrian Period. In 1946, Reg Sprigg noticed "jellyfishes" in the Ediacara Hills of Australia's Flinders Ranges, which were at the time believed to be Early Cambrian. It was not until the British discovery of the iconic Charnia that the Precambrian was seriously considered as containing life. This frond-shaped fossil was found in England's Charnwood Forest first by a 15 year-old girl in 1956 (Tina Negus, who was not believed) and then the next year by a group of three schoolboys including 15 year-old Roger Mason. Due to the detailed geological mapping of the British Geological Survey, there was no doubt these fossils sat in Precambrian rocks. Palaeontologist Martin Glaessner finally, in 1959, made the connection between this and the earlier finds and with a combination of improved dating of existing specimens and an injection of vigour into the search, many more instances were recognised. All specimens discovered until 1967 were in coarse-grained sandstone that prevented preservation of fine details, making interpretation difficult. S.B. Misra's discovery of fossiliferous ash-beds at the Mistaken Point assemblage in Newfoundland changed all this as the delicate detail preserved by the fine ash allowed the description of features that were previously undiscernible. It was also the first discovery of Ediacarans in deep water sediments. Poor communication, combined with the difficulty in correlating globally distinct formations, led to a plethora of different names for the biota. In 1960 the French name "Ediacarien" – after the Ediacara Hills – was added to the competing terms "Sinian" and "Vendian" for terminal-Precambrian rocks, and these names were also applied to the life-forms. "Ediacaran" and "Ediacarian" were subsequently applied to the epoch or period of geological time and its corresponding rocks. In March 2004, the International Union of Geological Sciences ended the inconsistency by formally naming the terminal period of the Neoproterozoic after the Australian locality. The term "Ediacaran biota" and similar ("Ediacara" / "Ediacaran" / "Ediacarian" / "Vendian" and "fauna" / "biota") has, at various times, been used in a geographic, stratigraphic, taphonomic, or biological sense, with the latter the most common in modern literature. ==Preservation==

Microbial mats -algal mat, salty lake on the White Sea seaside Microbial mats are areas of sediment stabilised by the presence of colonies of microbes that secrete sticky fluids or otherwise bind the sediment particles. They appear to migrate upwards when covered by a thin layer of sediment but this is an illusion caused by the colony's growth; individuals do not, themselves, move. If too thick a layer of sediment is deposited before they can grow or reproduce through it, parts of the colony will die leaving behind fossils with a characteristically wrinkled ("elephant skin") and tubercular texture. Some Ediacaran strata with the texture characteristics of microbial mats contain fossils, and Ediacaran fossils are almost always found in beds that contain these microbial mats. Although microbial mats were once widespread before the Cambrian substrate revolution, the evolution of grazing organisms vastly reduced their numbers. These communities are now limited to inhospitable refugia, such as the stromatolites found in Hamelin Pool Marine Nature Reserve in Shark Bay, Western Australia, where the salt levels can be twice those of the surrounding sea. Fossilization '' is barely distinguishable from the "elephant skin" texture on this cast. The preservation of Ediacaran fossils is of interest, since as soft-bodied organisms they would normally not fossilize. Further, unlike later soft-bodied fossil biota such as the Burgess Shale or Solnhofen Limestone, the Ediacaran biota is not found in a restricted environment subject to unusual local conditions: they are global. The processes that were operating must therefore have been systemic and worldwide. Something about the Ediacaran Period permitted these delicate creatures to be left behind; the fossils may have been preserved by virtue of rapid covering by ash or sand, trapping them against the mud or microbial mats on which they lived. Their preservation was possibly enhanced by the high concentration of silica in the oceans before silica-secreting organisms such as sponges and diatoms became prevalent. Ash beds provide more detail and can readily be dated to the nearest million years or better using radiometric dating. However, it is more common to find Ediacaran fossils under sandy beds deposited by storms or in turbidites formed by high-energy bottom-scraping ocean currents. Scale of preservation The rate of cementation of the overlying substrate relative to the rate of decomposition of the organism determines whether the top or bottom surface of an organism is preserved. Most disc-shaped fossils decomposed before the overlying sediment was cemented, whereupon ash or sand slumped in to fill the void, leaving a cast of the organism's underside. Conversely, quilted fossils tended to decompose after the cementation of the overlying sediment; hence their upper surfaces are preserved. Their more resistant nature is reflected in the fact that, in rare occasions, quilted fossils are found within storm beds as the high-energy sedimentation did not destroy them as it would have the less-resistant discs. Further, in some cases, the bacterial precipitation of minerals formed a "death mask", ultimately leaving a positive, cast-like impression of the organism. ==Morphology==

The Ediacaran biota exhibited a vast range of morphological characteristics. Size ranged from millimetres to metres; complexity from "blob-like" to intricate; rigidity from sturdy and resistant to jelly-soft. Almost all forms of symmetry were present. These organisms differed from earlier, mainly microbial, fossils in having an organised, differentiated multicellular construction and centimetre-plus sizes. generated intense media excitement though some have claimed they are instead inorganic structures formed by the precipitation of minerals on the inside of a hole. a view that, while it had enjoyed a notable gain of supporters as of 2007, has since suffered following further research comparing the potential Doushantuo embryos' morphologies with those of Thiomargarita specimens, both living and in various stages of decay. A recent discovery of comparable Ediacaran fossil embryos from the Portfjeld Formation in Greenland has significantly expanded the paleogeographical occurrence of Doushantuo-type fossil "embryos" with similar biotic forms now reported from differing paleolatitudes. : Microfossils dating from – just 3 million years after the end of the Cryogenian glaciations – may represent embryonic 'resting stages' in the life cycle of the earliest known animals. An alternative proposal is that these structures represent adult stages of the multicellular organisms of this period. Microfossils of Caveasphaera are thought to foreshadow the evolutionary origin of animal-like embryology. ; Discs : Circular fossils, such as Ediacaria, Cyclomedusa, and Rugoconites led to the initial identification of Ediacaran fossils as cnidaria, which include jellyfish and corals. and yet others may represent scratch marks formed as stalked organisms spun around their holdfasts. ; Bags : Fossils such as Pteridinium preserved within sediment layers resemble "mud-filled bags". The scientific community is a long way from reaching a consensus on their interpretation. ; Toroids : The fossil Vendoglossa tuberculata from the Nama Group, Namibia, has been interpreted as a dorso-ventrally compressed stem-group metazoan, with a large gut cavity and a transversely ridged ectoderm. The organism is in the shape of a flattened torus, with the long axis of its toroidal body running through the approximate center of the presumed gut cavity. ; Quilted organisms : The organisms considered in Seilacher's revised definition of the Vendobionta share a "quilted" appearance and resembled an inflatable mattress. Sometimes these quilts would be torn or ruptured prior to preservation: Such damaged specimens provide valuable clues in the reconstruction process. For example, the three (or more) petaloid fronds of Swartpuntia germsi could only be recognised in a posthumously damaged specimen – usually multiple fronds were hidden as burial squashed the organisms flat. These organisms appear to form two groups: the fractal rangeomorphs and the simpler erniettomorphs. Including such fossils as the iconic Charnia and Swartpuntia, the group is both the most iconic of the Ediacaran biota and the most difficult to place within the existing tree of life. Lacking any mouth, gut, reproductive organs, or indeed any evidence of internal anatomy, their lifestyle was somewhat peculiar by modern standards; the most widely accepted hypothesis holds that they sucked nutrients out of the surrounding seawater by osmotrophy or osmosis. However, others argue against this. ; Non-Vendobionts : Possible early forms of living phyla, excluding them from some definitions of the Ediacaran biota. The earliest such fossil is the reputed bilaterian Vernanimalcula claimed by some, however, to represent the infilling of an egg-sac or acritarch. In 2020, Ikaria wariootia was claimed to represent one of the oldest organisms with anterior and posterior differentiation. Later examples are almost universally accepted as bilaterians and include the mollusc-like Kimberella, Spriggina (pictured) and the shield-shaped Parvancorina, whose affinities are currently debated. A suite of fossils known as the small shelly fossils are represented in the Ediacaran, most famously by Cloudina, a shelly tube-like fossil that often shows evidence of predatory boring, suggesting that, while predation may not have been common in the Ediacaran Period, it was at least present. Organic microfossils known as small carbonaceous fossils are found in Ediacaran sediments, including the spiral-shaped Cochleatina which spans the Ediacaran–Cambrian boundary. Ediacaria also survived well into the Cambrian. Representatives of modern taxa existed in the Ediacaran, some of which are recognisable today. Sponges, red and green algæ, protists and bacteria are all easily recognisable with some pre-dating the Ediacaran by nearly three billion years. Possible arthropods have also been described. Surface trails left by Treptichnus bear similarities to modern priapulids. Fossils of the hard-shelled foraminifera Platysolenites are known from the latest Ediacaran of western Siberia, coexisting with Cloudina and Namacalathus. ; Filaments : Filament-shaped structures in Precambrian fossils have been observed on many occasions. Frondose fossils in Newfoundland have been observed to have developed filamentous bedding planes, inferred to be stolonic outgrowths. A study of Brazilian Ediacaran fossils found filamentous microfossils, suggested to be eukaryotes or large sulfur-oxidizing-bacteria (SOBs). Fungus-like filaments found in the Doushantuo Formation have been interpreted as eukaryotes and possibly fungi, providing possible evidence for the evolution and terrestrialization of fungi ~635 Ma. ; Trace fossils : With the exception of some very simple vertical burrows the only Ediacaran burrows are horizontal, lying on or just below the surface of the seafloor. Such burrows have been taken to imply the presence of motile organisms with heads, which would probably have had a bilateral symmetry. This could place them in the bilateral clade of animals but they could also have been made by simpler organisms feeding as they slowly rolled along the sea floor. Putative "burrows" dating as far back as may have been made by animals that fed on the undersides of microbial mats, which would have shielded them from a chemically unpleasant ocean; however their uneven width and tapering ends make a biological origin so difficult to defend that even the original proponent no longer believes they are authentic. : The burrows observed imply simple behaviour, and the complex efficient feeding traces common from the start of the Cambrian are absent. Some Ediacaran fossils, especially discs, have been interpreted tentatively as trace fossils but this hypothesis has not gained widespread acceptance. As well as burrows, some trace fossils have been found directly associated with an Ediacaran fossil. Yorgia and Dickinsonia are often found at the end of long pathways of trace fossils matching their shape; these fossils are thought to be associated with ciliary feeding but the precise method of formation of these disconnected and overlapping fossils largely remains a mystery. The potential mollusc Kimberella is associated with scratch marks, perhaps formed by a radula. ==Classification and interpretation==

in Chicago Classification of the Ediacarans is inevitably difficult, hence a variety of theories exist as to their placement on the tree of life. Martin Glaessner proposed in The Dawn of Animal Life (1984) that the Ediacaran biota were recognizable crown group members of modern phyla, but were unfamiliar because they had yet to evolve the characteristic features we use in modern classification. In 1998 Mark McMenamin claimed Ediacarans did not possess an embryonic stage, and thus could not be animals. He believed that they independently evolved a nervous system and brains, meaning that "the path toward intelligent life was embarked upon more than once on this planet". However, more recent discoveries have established that many of the circular forms formerly considered "cnidarian medusa" are actually holdfasts – sand-filled vesicles occurring at the base of the stem of upright frond-like Ediacarans. A notable example is the form known as Charniodiscus, a circular impression later found to be attached to the long 'stem' of a frond-like organism that now bears the name. The link between frond-like Ediacarans and sea pens has been thrown into doubt by multiple lines of evidence; chiefly the derived nature of the most frond-like pennatulacean octocorals, their absence from the fossil record before the Cenozoic, and the apparent cohesion between segments in Ediacaran frond-like organisms. Some researchers have suggested that an analysis of "growth poles" discredits the pennatulacean nature of Ediacaran fronds. Protozoans Adolf Seilacher has suggested that in the Ediacaran, animals take over from giant protists as the dominant life form. The modern xenophyophores are giant single-celled protozoans found throughout the world's oceans, largely on the abyssal plain. Genomic evidence suggests that the xenophyophores are a specialised group of Foraminifera. Unique phyla Seilacher has suggested that the Ediacaran organisms represented a unique and extinct grouping of related forms descended from a common ancestor (clade) and created the kingdom Vendozoa, named after the now-obsolete Vendian era. He later excluded fossils identified as metazoans and relaunched the phylum "Vendobionta", which he described as "quilted" cnidarians lacking stinging cells. This absence precludes the current cnidarian method of feeding, so Seilacher suggested that the organisms may have survived by symbiosis with photosynthetic or chemoautotrophic organisms. Mark McMenamin saw such feeding strategies as characteristic for the entire biota, and referred to the marine biota of this period as a "Garden of Ediacara". Lichen hypothesis -like wisps of ferruginized clay, and that Ediacaran fossils have been found in strata that he interprets as desert soils. The suggestion has been disputed by other scientists; some have described the evidence as ambiguous and unconvincing, for instance noting that Dickinsonia fossils have been found on rippled surfaces (suggesting a marine environment), while trace fossils like Radulichnus could not have been caused by needle ice as Retallack has proposed. Ben Waggoner notes that the suggestion would place the root of the Cnidaria back from around 900 mya to between 1500 mya and 2000 mya, contradicting much other evidence. Matthew Nelsen, examining phylogenies of ascomycete fungi and chlorophyte algae (components of lichens), calibrated for time, finds no support for the hypothesis that lichens predated the vascular plants. Other interpretations Several classifications have been used to accommodate the Ediacaran biota at some point, from algae, to protozoans, to fungi to bacterial or microbial colonies, to hypothetical intermediates between plants and animals. A new extant genus discovered in 2014, Dendrogramma, which at the time of discovery appeared to be a basal metazoan but of unknown taxonomic placement, had been noted to have similarities with the Ediacaran fauna. It has since been found to be a siphonophore, possibly even sections of a more complex species. == Origin ==

It took almost 4 billion years from the formation of the Earth for Ediacaran fossils to first appear, 655 million years ago. While putative fossils are reported from , the first uncontroversial evidence for life is found , and cells with nuclei certainly existed by . It could be that no special explanation is required: the slow process of evolution simply required 4 billion years to accumulate the necessary adaptations. Indeed, there does seem to be a slow increase in the maximum level of complexity seen over this time, with more and more complex forms of life evolving as time progresses, with traces of earlier semi-complex life such as Nimbia, found in the Twitya formation, and older rocks dating to in Kazakhstan. s might have delayed or prevented the establishment of multicellular life. On the early Earth, reactive elements, such as iron and uranium, existed in a reduced form that would react with any free oxygen produced by photosynthesising organisms. Oxygen would not be able to build up in the atmosphere until all the iron had rusted (producing banded iron formations), and all the other reactive elements had been oxidised. Donald Canfield detected records of the first significant quantities of atmospheric oxygen just before the first Ediacaran fossils appeared – and the presence of atmospheric oxygen was soon heralded as a possible trigger for the Ediacaran radiation. Oxygen seems to have accumulated in two pulses; the rise of small, sessile (stationary) organisms seems to correlate with an early oxygenation event, with larger and mobile organisms appearing around the second pulse of oxygenation. However, the assumptions underlying the reconstruction of atmospheric composition have attracted some criticism, with widespread anoxia having little effect on life where it occurs in the Early Cambrian and the Cretaceous. Periods of intense cold have also been suggested as a barrier to the evolution of multicellular life. The earliest known embryos, from China's Doushantuo Formation, appear just a million years after the Earth emerged from a global glaciation, suggesting that ice cover and cold oceans may have prevented the emergence of multicellular life. In early 2008, a team analysed the range of basic body structures ("disparity") of Ediacaran organisms from three different fossil beds: Avalon in Canada, to ; White Sea in Russia, to ; and Nama in Namibia, to , immediately before the start of the Cambrian. They found that, while the White Sea assemblage had the most species, there was no significant difference in disparity between the three groups, and concluded that before the beginning of the Avalon timespan these organisms must have gone through their own evolutionary "explosion", which may have been similar to the famous Cambrian explosion. Preservation bias The paucity of Ediacaran fossils after the Cambrian could simply be due to conditions no longer favoring the fossilization of Ediacaran organisms, which may have continued to thrive unpreserved for a considerable time. might be expected in exceptionally preserved fossil assemblages (Konservat-Lagerstätten) such as the Burgess Shale and Chengjiang. Although no reports of Ediacara-type organisms in the Cambrian period are widely accepted at present, a few disputed reports have been made, as well as unpublished observations of 'vendobiont' fossils from 535 Ma Orsten-type deposits in China. Predation and grazing '' might have had a predatory or grazing lifestyle. It has been suggested that by the Early Cambrian, organisms higher in the food chain caused the microbial mats to largely disappear. If these grazers first appeared as the Ediacaran biota started to decline, then it may suggest that they destabilised the microbial mats in a "Cambrian substrate revolution", leading to displacement or detachment of the biota; or that the destruction of the microbial substrate destabilized the ecosystem, causing extinctions. Alternatively, skeletonized animals could have fed directly on the relatively undefended Ediacaran biota. However, if the interpretation of the Ediacaran age Kimberella as a grazer is correct then this suggests that the biota had already had limited exposure to "predation". However, the supposed "competitive exclusion" of brachiopods by bivalve molluscs was eventually deemed to be a coincidental result of two unrelated trends. Change in environmental conditions Great changes were happening at the end of the Precambrian and the start of the Early Cambrian. The breakup of the supercontinents, rising sea levels (creating shallow, "life-friendly" seas), a nutrient crisis, fluctuations in atmospheric composition, including oxygen and carbon dioxide levels, and changes in ocean chemistry (promoting biomineralisation) could all have played a part. ==Assemblages==

Late Ediacaran macrofossils are recognized globally in at least 52 formations and a variety of depositional conditions. Each formation is commonly grouped into three main types, known as assemblages and named after typical localities. Each assemblage tends to occupy its own time period and region of morphospace, and after an initial burst of diversification (or extinction) changes little for the rest of its existence. The assemblage is easily dated because it contains many fine ash-beds, which are a good source of zircons used in the uranium-lead method of radiometric dating. These fine-grained ash beds also preserve exquisite detail. Constituents of this biota appear to survive through until the extinction of all Ediacarans at the base of the Cambrian. One interpretation of the biota is as deep-sea-dwelling rangeomorphs such as Charnia, all of which share a fractal growth pattern. They were probably preserved in situ (without post-mortem transportation), although this point is not universally accepted. The assemblage, while less diverse than the White Sea or Nama assemblages, resembles Carboniferous suspension-feeding communities, which may suggest filter feeding as the assemblage is often found in water too deep for photosynthesis. White Sea assemblage The White Sea or Ediacaran assemblage is named after Russia's White Sea or Australia's Ediacara Hills and is marked by much higher diversity than the Avalon or Nama assemblages. but a few are preserved within sandy units. Nama assemblage The Nama assemblage is best represented in Namibia. It is marked by extreme biotic turnover, with rates of extinction exceeding rates of origination for the whole period. These beds are sandwiched between units comprising interbedded sandstones, siltstones and shales—with microbial mats, where present, usually containing the fossils. The environment is interpreted as sand bars formed at the mouth of a delta's distributaries. Significance of assemblages Since they are globally distributed – described on all continents except Antarctica – geographical boundaries do not appear to be a factor; the same fossils are found at all palaeolatitudes (the latitude where the fossil was created, accounting for continental drift - an application of paleomagnetism) and in separate sedimentary basins. Because of this, the three assemblages are often separated by temporal boundaries rather than environmental ones (timeline at right). As the Ediacaran biota represent an early stage in multicellular life's history, it is unsurprising that not all possible modes of life are occupied. It has been estimated that of 92 potentially possible modes of life – combinations of feeding style, tiering and motility — no more than a dozen are occupied by the end of the Ediacaran. Just four are represented in the Avalon assemblage. ==See also==