Vendian biota
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The Ediacaran (; formerly Vendian) biota is a taxonomic period classification that consists of all life forms that were present on Earth during the
Ediacaran The Ediacaran ( ) is a geological period of the Neoproterozoic geologic era, Era that spans 96 million years from the end of the Cryogenian Period at 635 Million years ago, Mya to the beginning of the Cambrian Period at 538.8 Mya. It is the last ...
Period (). These were enigmatic tubular and frond-shaped, mostly sessile, organisms.
Trace fossil A trace fossil, also called an ichnofossil (; ), is a fossil record of biological activity by lifeforms, but not the preserved remains of the organism itself. Trace fossils contrast with body fossils, which are the fossilized remains of part ...
s of these organisms have been found worldwide, and represent the earliest known complex
multicellular organism A multicellular organism is an organism that consists of more than one cell (biology), cell, unlike unicellular organisms. All species of animals, Embryophyte, land plants and most fungi are multicellular, as are many algae, whereas a few organism ...
s. The term "Ediacara biota" has received criticism from some scientists due to its alleged inconsistency, arbitrary exclusion of certain fossils, and inability to be precisely defined. The Ediacaran biota may have undergone
evolutionary radiation An evolutionary radiation is an increase in taxonomic diversity that is caused by elevated rates of speciation, that may or may not be associated with an increase in morphological disparity. A significantly large and diverse radiation within ...
in a proposed event called the Avalon explosion, . This was after the Earth had thawed from the
Cryogenian The Cryogenian (from , meaning "cold" and , romanized: , meaning "birth") is a geologic period that lasted from . It is the second of the three periods of the Neoproterozoic era, preceded by the Tonian and followed by the Ediacaran. The Cryoge ...
period's extensive glaciation. This biota largely disappeared with the rapid increase in
biodiversity Biodiversity is the variability of life, life on Earth. It can be measured on various levels. There is for example genetic variability, species diversity, ecosystem diversity and Phylogenetics, phylogenetic diversity. Diversity is not distribut ...
known as the Cambrian explosion. Most of the currently existing
body plan A body plan, (), or ground plan is a set of morphology (biology), morphological phenotypic trait, features common to many members of a phylum of animals. The vertebrates share one body plan, while invertebrates have many. This term, usually app ...
s of
animal Animals are multicellular, eukaryotic organisms in the Biology, biological Kingdom (biology), kingdom Animalia (). With few exceptions, animals heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, ...
s first appeared in the
fossil record A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
of the
Cambrian The Cambrian ( ) is the first geological period of the Paleozoic Era, and the Phanerozoic Eon. The Cambrian lasted 51.95 million years from the end of the preceding Ediacaran period 538.8 Ma (million years ago) to the beginning of the Ordov ...
rather than the Ediacaran. For macroorganisms, the Cambrian biota appears to have almost completely replaced the organisms that dominated the Ediacaran fossil record, although relationships are still a matter of debate. The organisms of the Ediacaran Period first appeared around and flourished until the cusp of the
Cambrian The Cambrian ( ) is the first geological period of the Paleozoic Era, and the Phanerozoic Eon. The Cambrian lasted 51.95 million years from the end of the preceding Ediacaran period 538.8 Ma (million years ago) to the beginning of the Ordov ...
, when the characteristic communities of fossils vanished. A diverse Ediacaran community was discovered in 1995 in
Sonora Sonora (), officially Estado Libre y Soberano de Sonora (), is one of the 31 states which, along with Mexico City, comprise the Administrative divisions of Mexico, Federal Entities of Mexico. The state is divided into Municipalities of Sonora, 72 ...
, Mexico, and is approximately 555 million years in age, roughly coeval with Ediacaran fossils of the
Ediacara Hills Ediacara Hills ( ), also known as Ediacaran Hills, are a range of low hills in the northern part of the Flinders Ranges of South Australia, around north of the state capital of Adelaide city centre, Adelaide. They are within the Nilpena Ediaca ...
in
South Australia South Australia (commonly abbreviated as SA) is a States and territories of Australia, state in the southern central part of Australia. With a total land area of , it is the fourth-largest of Australia's states and territories by area, which in ...
and the
White Sea The White Sea (; Karelian language, Karelian and ; ) is a southern inlet of the Barents Sea located on the northwest coast of Russia. It is surrounded by Karelia to the west, the Kola Peninsula to the north, and the Kanin Peninsula to the nort ...
on the coast of
Russia Russia, or the Russian Federation, is a country spanning Eastern Europe and North Asia. It is the list of countries and dependencies by area, largest country in the world, and extends across Time in Russia, eleven time zones, sharing Borders ...
. While rare fossils that may represent survivors have been found as late as the
Middle Cambrian Middle or The Middle may refer to: * Centre (geometry), the point equally distant from the outer limits. Places * Middle (sheading), a subdivision of the Isle of Man * Middle Bay (disambiguation) * Middle Brook (disambiguation) * Middle Creek (di ...
(510–500 Mya), the earlier fossil communities disappear from the record at the end of the Ediacaran leaving only curious fragments of once-thriving
ecosystem An ecosystem (or ecological system) is a system formed by Organism, organisms in interaction with their Biophysical environment, environment. The Biotic material, biotic and abiotic components are linked together through nutrient cycles and en ...
s. Multiple hypotheses exist to explain the disappearance of this biota, including
preservation bias Taphonomy is the study of how organisms decay and become fossilized or preserved in the paleontological record. The term ''taphonomy'' (from Greek , 'burial' and , 'law') was introduced to paleontology in 1940 by Soviet scientist Ivan Efremov ...
, a changing environment, the advent of predators and competition from other life-forms. A sampling, reported in 2018, of late Ediacaran strata across the scattered remnants of
Baltica Baltica is a paleocontinent that formed in the Paleoproterozoic and now constitutes northwestern Eurasia, or Europe north of the Trans-European Suture Zone and west of the Ural Mountains. The thick core of Baltica, the East European Craton, i ...
suggests the flourishing of the organisms coincided with conditions of low overall productivity with a very high percentage produced by bacteria, which may have led to high concentrations of dissolved organic material in the oceans. Determining where Ediacaran organisms fit in the
tree of life The tree of life is a fundamental archetype in many of the world's mythology, mythological, religion, religious, and philosophy, philosophical traditions. It is closely related to the concept of the sacred tree.Giovino, Mariana (2007). ''The ...
has proven challenging; it is not even established that most of them were animals, with suggestions that they were
lichens A lichen ( , ) is a hybrid colony (biology), colony of algae or cyanobacteria living symbiotically among hypha, filaments of multiple fungus species, along with yeasts and bacteria embedded in the cortex or "skin", in a mutualism (biology), m ...
(fungus-alga symbionts),
algae Algae ( , ; : alga ) is an informal term for any organisms of a large and diverse group of photosynthesis, photosynthetic organisms that are not plants, and includes species from multiple distinct clades. Such organisms range from unicellular ...
,
protists A protist ( ) or protoctist is any Eukaryote, eukaryotic organism that is not an animal, Embryophyte, land plant, or fungus. Protists do not form a Clade, natural group, or clade, but are a Paraphyly, paraphyletic grouping of all descendants o ...
known as
foraminifera Foraminifera ( ; Latin for "hole bearers"; informally called "forams") are unicellular organism, single-celled organisms, members of a phylum or class (biology), class of Rhizarian protists characterized by streaming granular Ectoplasm (cell bio ...
,
fungi A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
or
microbial A microorganism, or microbe, is an organism of microscopic size, which may exist in its single-celled form or as a colony of cells. The possible existence of unseen microbial life was suspected from antiquity, with an early attestation in ...
colonies, or hypothetical intermediates between plants and animals. The morphology and habit of some taxa (e.g. '' Funisia dorothea'') suggest relationships to
Porifera Sponges or sea sponges are primarily marine invertebrates of the animal phylum Porifera (; meaning 'pore bearer'), a Basal (phylogenetics) , basal clade and a sister taxon of the Eumetazoa , diploblasts. They are sessility (motility) , sessile ...
or
Cnidaria Cnidaria ( ) is a phylum under kingdom Animalia containing over 11,000 species of aquatic invertebrates found both in fresh water, freshwater and marine environments (predominantly the latter), including jellyfish, hydroid (zoology), hydroids, ...
(e.g. '' Auroralumina''). ''
Kimberella ''Kimberella'' is an extinct genus of marine bilaterian known only from rocks of the Ediacaran period. The slug-like organism fed by scratching the microbial surface on which it dwelt in a manner similar to the gastropods, although its affinity ...
'' may show a similarity to
mollusc Mollusca is a phylum of protostome, protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant taxon, extant species of molluscs are recognized, making it the second-largest animal phylum ...
s, and other organisms have been thought to possess
bilateral symmetry Symmetry in biology refers to the symmetry observed in organisms, including plants, animals, fungi, and bacteria. External symmetry can be easily seen by just looking at an organism. For example, the face of a human being has a plane of symme ...
, although this is controversial. Most macroscopic fossils are morphologically distinct from later life-forms: they resemble discs, tubes, mud-filled bags or quilted mattresses. Due to the difficulty of deducing evolutionary relationships among these organisms, some
palaeontologist Paleontology, also spelled as palaeontology or palæontology, is the scientific study of the life of the past, mainly but not exclusively through the study of fossils. Paleontologists use fossils as a means to classify organisms, measure geolo ...
s have suggested that these represent completely extinct lineages that do not resemble any living organism. Palaeontologist Adolf Seilacher proposed a separate subkingdom level category Vendozoa (now renamed Vendobionta) in the Linnaean hierarchy for the Ediacaran biota. If these enigmatic organisms left no descendants, their strange forms might be seen as a "failed experiment" in multicellular life, with later multicellular life evolving independently from unrelated single-celled organisms. A 2018 study confirmed that one of the period's most-prominent and iconic fossils, ''
Dickinsonia ''Dickinsonia'' is a genus of extinct organism that lived during the late Ediacaran period in what is now Australia, China, Russia, and Ukraine. It had a round, bilaterally symmetric body with multiple segments running along it. It could range f ...
'', included
cholesterol Cholesterol is the principal sterol of all higher animals, distributed in body Tissue (biology), tissues, especially the brain and spinal cord, and in Animal fat, animal fats and oils. Cholesterol is biosynthesis, biosynthesized by all anima ...
, suggesting affinities to animals, fungi, or red algae.


History

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 Newfoundland and Labrador is the easternmost province of Canada, in the country's Atlantic region. The province comprises the island of Newfoundland and the continental region of Labrador, having a total size of . As of 2025 the population ...
. However, since they lay below the "Primordial Strata" of the
Cambrian The Cambrian ( ) is the first geological period of the Paleozoic Era, and the Phanerozoic Eon. The Cambrian lasted 51.95 million years from the end of the preceding Ediacaran period 538.8 Ma (million years ago) to the beginning of the Ordov ...
that was then thought to contain the very first signs of animal life, a proposal four years after their discovery by Elkanah Billings that these simple forms represented fauna was dismissed by his peers. Instead, they were interpreted as gas escape structures or inorganic
concretion A concretion is a hard and compact mass formed by the precipitation of mineral cement within the spaces between particles, and is found in sedimentary rock or soil. Concretions are often ovoid or spherical in shape, although irregular shapes a ...
s. No similar structures elsewhere in the world were then known and the one-sided debate soon fell into obscurity. In 1933, Georg Gürich discovered specimens in
Namibia Namibia, officially the Republic of Namibia, is a country on the west coast of Southern Africa. Its borders include the Atlantic Ocean to the west, Angola and Zambia to the north, Botswana to the east and South Africa to the south; in the no ...
but assigned them to the Cambrian Period. In 1946, Reg Sprigg noticed "jellyfishes" in the
Ediacara Hills Ediacara Hills ( ), also known as Ediacaran Hills, are a range of low hills in the northern part of the Flinders Ranges of South Australia, around north of the state capital of Adelaide city centre, Adelaide. They are within the Nilpena Ediaca ...
of Australia's
Flinders Ranges The Flinders Ranges are the largest mountain ranges in South Australia, which starts about north of Adelaide. The ranges stretch for over from Port Pirie to Lake Callabonna. The Adnyamathanha people are the Aboriginal group who have inhab ...
, which were at the time believed to be Early Cambrian. It was not until the British discovery of the iconic '' Charnia'' that the
Precambrian The Precambrian ( ; or pre-Cambrian, sometimes abbreviated pC, or Cryptozoic) is the earliest part of Earth's history, set before the current Phanerozoic Eon. The Precambrian is so named because it preceded the Cambrian, the first period of t ...
was seriously considered as containing life. This
frond A frond is a large, divided leaf. In both common usage and botanical nomenclature, the leaves of ferns are referred to as fronds and some botanists restrict the term to this group. Other botanists allow the term frond to also apply to the lar ...
-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 map A geological map or geologic map is a special-purpose map made to show various geological features. Rock (geology), Rock units or stratum, geologic strata are shown by color or symbols. Bed (geology), Bedding planes and structural features such ...
ping of the
British Geological Survey The British Geological Survey (BGS) is a partly publicly funded body which aims to advance Earth science, geoscientific knowledge of the United Kingdom landmass and its continental shelf by means of systematic surveying, monitoring and research. ...
, 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 Sandstone is a Clastic rock#Sedimentary clastic rocks, clastic sedimentary rock composed mainly of grain size, sand-sized (0.0625 to 2 mm) silicate mineral, silicate grains, Cementation (geology), cemented together by another mineral. Sand ...
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 The geologic time scale or geological time scale (GTS) is a representation of time based on the rock record of Earth. It is a system of chronological dating that uses chronostratigraphy (the process of relating strata to time) and geochronolo ...
and its corresponding rocks. In March 2004, the
International Union of Geological Sciences The International Union of Geological Sciences (IUGS) is an international non-governmental organization devoted to global cooperation in the field of geology. As of 2023, it represents more than 1 million geoscientists around the world. About Fo ...
ended the inconsistency by formally naming the terminal period of the
Neoproterozoic The Neoproterozoic Era is the last of the three geologic eras of the Proterozoic geologic eon, eon, spanning from 1 billion to 538.8 million years ago, and is the last era of the Precambrian "supereon". It is preceded by the Mesoproterozoic era an ...
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

Microbial mat A microbial mat is a multi-layered sheet or biofilm of microbial colonies, composed of mainly bacteria and/or archaea. Microbial mats grow at interfaces between different types of material, mostly on submerged or moist surfaces, but a few surviv ...
s 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 Stromatolites ( ) or stromatoliths () are layered sedimentary formations ( microbialite) that are created mainly by photosynthetic microorganisms such as cyanobacteria, sulfate-reducing bacteria, and Pseudomonadota (formerly proteobacteria) ...
found in Hamelin Pool Marine Nature Reserve in
Shark Bay Shark Bay () is a World Heritage Site in the Gascoyne region of Western Australia. The area is located approximately north of Perth, on the westernmost point of the Australian continent. UNESCO's listing of Shark Bay as a World Heritage S ...
,
Western Australia Western Australia (WA) is the westernmost state of Australia. It is bounded by the Indian Ocean to the north and west, the Southern Ocean to the south, the Northern Territory to the north-east, and South Australia to the south-east. Western Aust ...
, where the salt levels can be twice those of the surrounding sea.


Fossilization

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 The Burgess Shale is a fossil-bearing deposit exposed in the Canadian Rockies of British Columbia, Canada. It is famous for the exceptional preservation of the soft parts of its fossils. At old (middle Cambrian), it is one of the earliest fos ...
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 Radiometric dating, radioactive dating or radioisotope dating is a technique which is used to Chronological dating, date materials such as Rock (geology), rocks or carbon, in which trace radioactive impurity, impurities were selectively incorporat ...
. However, it is more common to find Ediacaran fossils under sandy beds deposited by storms or in
turbidite A turbidite is the geologic Deposition (geology), deposit of a turbidity current, which is a type of amalgamation of fluidal and sediment gravity flow responsible for distributing vast amounts of clastic sediment into the deep ocean. Sequencing ...
s formed by high-energy bottom-scraping ocean currents. Soft-bodied organisms today rarely fossilize during such events, but the presence of widespread microbial mats probably aided preservation by stabilising their impressions in the sediment below.


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
bacteria Bacteria (; : bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of Prokaryote, prokaryotic microorganisms. Typically a few micr ...
l
precipitation In meteorology, precipitation is any product of the condensation of atmospheric water vapor that falls from clouds due to gravitational pull. The main forms of precipitation include drizzle, rain, rain and snow mixed ("sleet" in Commonwe ...
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 Symmetry () in everyday life refers to a sense of harmonious and beautiful proportion and balance. In mathematics, the term has a more precise definition and is usually used to refer to an object that is Invariant (mathematics), invariant und ...
were present. These organisms differed from earlier, mainly microbial, fossils in having an organised, differentiated multicellular construction and centimetre-plus sizes. These disparate morphologies can be broadly grouped into form taxa: ; "Embryos" : Recent discoveries of Precambrian multicellular life have been dominated by reports of embryos, particularly from the Doushantuo Formation in China. Some finds 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. Other "embryos" have been interpreted as the remains of the giant
sulfur Sulfur ( American spelling and the preferred IUPAC name) or sulphur ( Commonwealth spelling) is a chemical element; it has symbol S and atomic number 16. It is abundant, multivalent and nonmetallic. Under normal conditions, sulfur atoms ...
-reducing bacteria akin to '' Thiomargarita'',e.g. , summarised by a view that, while it had enjoyed a notable gain of supportersXiao ''et al.''.'s response to Bailey ''et al.''.'s original paper :
And Bailey ''et al.''.'s reply:
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 ''Rugoconites'' is a genus of Ediacaran biota found as fossils in the form of a circular or oval-like impression preserved in high relief, six or more centimeters in diameter. The fossils are surrounded by frills that have been interpreted ( Wade ...
'' led to the initial identification of Ediacaran fossils as
cnidaria Cnidaria ( ) is a phylum under kingdom Animalia containing over 11,000 species of aquatic invertebrates found both in fresh water, freshwater and marine environments (predominantly the latter), including jellyfish, hydroid (zoology), hydroids, ...
, which include jellyfish and corals. Further examination has provided alternative interpretations of all disc-shaped fossils: not one is now confidently recognised as a jellyfish. Alternate explanations include holdfasts and
protist A protist ( ) or protoctist is any eukaryotic organism that is not an animal, land plant, or fungus. Protists do not form a natural group, or clade, but are a paraphyletic grouping of all descendants of the last eukaryotic common ancest ...
s; the patterns displayed where two meet have led to many 'individuals' being identified as microbial colonies, 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.(a) The only current description, far from universal acceptance, appears as: ; 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 ectoderm is one of the three primary germ layers formed in early embryonic development. It is the outermost layer, and is superficial to the mesoderm (the middle layer) and endoderm (the innermost layer). It emerges and originates from the o ...
. 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 A mattress is a large, usually rectangular pad for supporting a person Lying (position), lying down, especially for sleeping. It is designed to be used as a bed, or on a bed frame as part of a bed. Mattresses may consist of a Quilting, quilted o ...
. 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 In mathematics, a fractal is a Shape, geometric shape containing detailed structure at arbitrarily small scales, usually having a fractal dimension strictly exceeding the topological dimension. Many fractals appear similar at various scale ...
rangeomorph The rangeomorphs are a group of Ediacaran Ediacaran biota, fossils. Ediacarans are the oldest large fossil organisms on earth, and many are not self-evidently related to anything else that has ever lived. However, some Ediacarans clearly resemble ...
s 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 The tree of life is a fundamental archetype in many of the world's mythology, mythological, religion, religious, and philosophy, philosophical traditions. It is closely related to the concept of the sacred tree.Giovino, Mariana (2007). ''The ...
. 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 Osmosis (, ) is the spontaneous net movement or diffusion of solvent molecules through a selectively permeable membrane, selectively-permeable membrane from a region of high water potential (region of lower solute concentration) to a region of ...
. However, others argue against this. ; Non-Vendobionts : Possible early forms of
living Living or The Living may refer to: Common meanings *Life, a condition that distinguishes organisms from inorganic objects and dead organisms ** Living species, one that is not extinct *Personal life, the course of an individual human's life * ...
phyla Phyla, the plural of ''phylum'', may refer to: * Phylum, a biological taxon between Kingdom and Class * by analogy, in linguistics, a large division of possibly related languages, or a major language family which is not subordinate to another Phy ...
, excluding them from some definitions of the Ediacaran biota. The earliest such fossil is the reputed bilaterian ''
Vernanimalcula ''Vernanimalcula guizhouena'' is an acritarch dating from ; it was between 0.1 and 0.2 mm across (roughly the width of one or two human hairs). ''Vernanimalcula'' means "small spring animal", referring to its appearance in the fossil record ...
'' claimed by some, however, to represent the infilling of an egg-sac or
acritarch Acritarchs () are organic microfossils, known from the Precambrian to the present. The classification is a catch all term used to refer to any organic microfossils that cannot be assigned to other groups. Their diversity reflects major ecolog ...
. In 2020, ''
Ikaria wariootia ''Ikaria wariootia'' is an early example of a wormlike, bilaterian organism. Its fossils are found in rocks of the Ediacara Member of South Australia that are estimated to be between 560 and 555 million years old. A representative of the Ediaca ...
'' 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 ''Kimberella'' is an extinct genus of marine bilaterian known only from rocks of the Ediacaran period. The slug-like organism fed by scratching the microbial surface on which it dwelt in a manner similar to the gastropods, although its affinity ...
'', ''
Spriggina ''Spriggina'' is a genus of early animals whose relationship to living animals is unclear. Fossils of ''Spriggina'' are known from the late Ediacaran period in what is now South Australia. ''Spriggina floundersi'' is the official fossil emblem o ...
'' (pictured) and the shield-shaped '' Parvancorina'' whose affinities are currently debated. A suite of fossils known as the
small shelly fossil The small shelly fauna, small shelly fossils (SSF), or early skeletal fossils (ESF) are biomineralization, mineralized fossils, many only a few millimetres long, with a nearly continuous record from the latest stages of the Ediacaran to the end o ...
s 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.
Sponge Sponges or sea sponges are primarily marine invertebrates of the animal phylum Porifera (; meaning 'pore bearer'), a basal clade and a sister taxon of the diploblasts. They are sessile filter feeders that are bound to the seabed, and a ...
s, red and green
algæ Algae ( , ; : alga ) is an informal term for any organisms of a large and diverse group of photosynthetic organisms that are not plants, and includes species from multiple distinct clades. Such organisms range from unicellular microalgae, suc ...
,
protist A protist ( ) or protoctist is any eukaryotic organism that is not an animal, land plant, or fungus. Protists do not form a natural group, or clade, but are a paraphyletic grouping of all descendants of the last eukaryotic common ancest ...
s and
bacteria Bacteria (; : bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of Prokaryote, prokaryotic microorganisms. Typically a few micr ...
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 Foraminifera ( ; Latin for "hole bearers"; informally called "forams") are unicellular organism, single-celled organisms, members of a phylum or class (biology), class of Rhizarian protists characterized by streaming granular Ectoplasm (cell bio ...
'' Platysolenites'' are known from the latest Ediacaran of western
Siberia Siberia ( ; , ) is an extensive geographical region comprising all of North Asia, from the Ural Mountains in the west to the Pacific Ocean in the east. It has formed a part of the sovereign territory of Russia and its predecessor states ...
, coexisting with ''Cloudina'' and ''
Namacalathus ''Namacalathus'' is a problematic metazoan fossil occurring in the latest Ediacaran. The first, and only described species, ''N. hermanastes'', was first described in 2000 from the Nama Group of central and southern Namibia. Discovery and nami ...
''. ; Filaments : Filament-shaped structures in Precambrian fossils have been observed on many occasions.
Frondose Frondosity (from Latin ''frondōsus'' meaning 'leafy') is the property of an organism that normally flourishes with fronds or leaf-like structures. Many frondose organisms are thalloid and lack the organization of tissues into organs, with the ...
fossils in Newfoundland have been observed to have developed filamentous bedding planes, inferred to be
stolon In biology, a stolon ( from Latin ''wikt:stolo, stolō'', genitive ''stolōnis'' – "branch"), also known as a runner, is a horizontal connection between parts of an organism. It may be part of the organism, or of its skeleton. Typically, animal ...
ic 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 fossil A trace fossil, also called an ichnofossil (; ), is a fossil record of biological activity by lifeforms, but not the preserved remains of the organism itself. Trace fossils contrast with body fossils, which are the fossilized remains of part ...
s : 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 In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
of
animal Animals are multicellular, eukaryotic organisms in the Biology, biological Kingdom (biology), kingdom Animalia (). With few exceptions, animals heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, ...
s 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 ''Dickinsonia'' is a genus of extinct organism that lived during the late Ediacaran period in what is now Australia, China, Russia, and Ukraine. It had a round, bilaterally symmetric body with multiple segments running along it. It could range f ...
'' 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 Mollusca is a phylum of protostome, protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant taxon, extant species of molluscs are recognized, making it the second-largest animal phylum ...
''Kimberella'' is associated with scratch marks, perhaps formed by a
radula The radula (; : radulae or radulas) is an anatomical structure used by mollusks for feeding, sometimes compared to a tongue. It is a minutely toothed, chitinous ribbon, which is typically used for scraping or cutting food before the food enters ...
.According to
: For a more cynical perspective see


Classification and interpretation

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 In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor ...
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
embryo An embryo ( ) is the initial stage of development for a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sp ...
nic stage, and thus could not be animals. He believed that they independently evolved a
nervous system In biology, the nervous system is the complex system, highly complex part of an animal that coordinates its behavior, actions and sense, sensory information by transmitting action potential, signals to and from different parts of its body. Th ...
and
brain The brain is an organ (biology), organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It consists of nervous tissue and is typically located in the head (cephalization), usually near organs for ...
s, meaning that "the path toward intelligent life was embarked upon more than once on this planet". In 2018 analysis of ancient
sterol A sterol is any organic compound with a Skeletal formula, skeleton closely related to Cholestanol, cholestan-3-ol. The simplest sterol is gonan-3-ol, which has a formula of , and is derived from that of gonane by replacement of a hydrogen atom on ...
s was taken as evidence that one of the period's most-prominent and iconic fossils, ''
Dickinsonia ''Dickinsonia'' is a genus of extinct organism that lived during the late Ediacaran period in what is now Australia, China, Russia, and Ukraine. It had a round, bilaterally symmetric body with multiple segments running along it. It could range f ...
'', was an early animal.


Cnidarians

Since the most primitive eumetazoans—multi-cellular animals with tissues—are
cnidarians Cnidaria ( ) is a phylum under kingdom Animalia containing over 11,000 species of aquatic invertebrates found both in fresh water, freshwater and marine environments (predominantly the latter), including jellyfish, hydroid (zoology), hydroids, ...
, and the first recognized Ediacaran fossil '' Charnia'' looks very much like a
sea pen Sea pens are marine cnidarians belonging to the superfamily Pennatuloidea, which are colony-forming benthic filter feeders within the order Scleralcyonacea. There are 14 families within the order and 35 extant genera, and it is estimated a ...
, the first attempt to categorise these fossils designated them as
jellyfish Jellyfish, also known as sea jellies or simply jellies, are the #Life cycle, medusa-phase of certain gelatinous members of the subphylum Medusozoa, which is a major part of the phylum Cnidaria. Jellyfish are mainly free-swimming marine animal ...
and
sea pen Sea pens are marine cnidarians belonging to the superfamily Pennatuloidea, which are colony-forming benthic filter feeders within the order Scleralcyonacea. There are 14 families within the order and 35 extant genera, and it is estimated a ...
s. 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 Tertiary, 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
protist A protist ( ) or protoctist is any eukaryotic organism that is not an animal, land plant, or fungus. Protists do not form a natural group, or clade, but are a paraphyletic grouping of all descendants of the last eukaryotic common ancest ...
s as the dominant life form. The modern xenophyophores are giant single-celled protozoans found throughout the world's oceans, largely on the
abyssal plain An abyssal plain is an underwater plain on the deep ocean floor, usually found at depths between . Lying generally between the foot of a continental rise and a mid-ocean ridge, abyssal plains cover more than 50% of the Earth's surface. They ...
. Genomic evidence suggests that the xenophyophores are a specialised group of
Foraminifera Foraminifera ( ; Latin for "hole bearers"; informally called "forams") are unicellular organism, single-celled organisms, members of a phylum or class (biology), class of Rhizarian protists characterized by streaming granular Ectoplasm (cell bio ...
.


Unique phyla

Seilacher has suggested that the Ediacaran organisms represented a unique and extinct grouping of related forms descended from a common ancestor (
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
) and created the kingdom Vendozoa, named after the now-obsolete
Vendian The Ediacaran ( ) is a geological period of the Neoproterozoic geologic era, Era that spans 96 million years from the end of the Cryogenian Period at 635 Million years ago, Mya to the beginning of the Cambrian Period at 538.8 Mya. It is the last ...
era. He later excluded fossils identified as
metazoa Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia (). With few exceptions, animals consume organic material, breathe oxygen, have myocytes and are able to move, can reproduce sexually, and grow from a hol ...
ns and relaunched the
phylum In biology, a phylum (; : phyla) is a level of classification, or taxonomic rank, that is below Kingdom (biology), kingdom and above Class (biology), class. Traditionally, in botany the term division (taxonomy), division has been used instead ...
"Vendobionta", which he described as "quilted"
cnidarians Cnidaria ( ) is a phylum under kingdom Animalia containing over 11,000 species of aquatic invertebrates found both in fresh water, freshwater and marine environments (predominantly the latter), including jellyfish, hydroid (zoology), hydroids, ...
lacking stinging cells. This absence precludes the current cnidarian method of feeding, so Seilacher suggested that the organisms may have survived by
symbiosis Symbiosis (Ancient Greek : living with, companionship < : together; and ''bíōsis'': living) is any type of a close and long-term biological interaction, between two organisms of different species. The two organisms, termed symbionts, can fo ...
with
photosynthetic Photosynthesis ( ) is a Biological system, system of biological processes by which Photoautotrophism, photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical ener ...
or
chemoautotrophic A chemotroph is an organism that obtains energy by the oxidation of electron donors in their environments. These molecules can be organic (chemoorganotrophs) or inorganic (chemolithotrophs). The chemotroph designation is in contrast to phototroph ...
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

Greg Retallack has proposed that Ediacaran organisms were
lichen A lichen ( , ) is a hybrid colony (biology), colony of algae or cyanobacteria living symbiotically among hypha, filaments of multiple fungus species, along with yeasts and bacteria embedded in the cortex or "skin", in a mutualism (biology), m ...
s. He argues that thin sections of Ediacaran fossils show lichen-like compartments and
hypha A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one o ...
-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 Cnidaria ( ) is a phylum under kingdom Animalia containing over 11,000 species of aquatic invertebrates found both in fresh water, freshwater and marine environments (predominantly the latter), including jellyfish, hydroid (zoology), hydroids, ...
back from around 900 mya to between 1500 mya and 2000 mya, contradicting much other evidence. Matthew Nelsen, examining phylogenies of
ascomycete Ascomycota is a phylum of the kingdom Fungi that, together with the Basidiomycota, forms the subkingdom Dikarya. Its members are commonly known as the sac fungi or ascomycetes. It is the largest phylum of Fungi, with over 64,000 species. The def ...
fungi and
chlorophyte Chlorophyta is a division (botany), division of green algae informally called chlorophytes. Description Chlorophytes are eukaryotic organisms composed of cells with a variety of coverings or walls, and usually a single green chloroplast in ea ...
algae (components of lichens), calibrated for time, finds no support for the hypothesis that lichens predated the
vascular plant Vascular plants (), also called tracheophytes (, ) or collectively tracheophyta (; ), are plants that have lignin, lignified tissues (the xylem) for conducting water and minerals throughout the plant. They also have a specialized non-lignified Ti ...
s.


Other interpretations

Several classifications have been used to accommodate the Ediacaran biota at some point, from
algae Algae ( , ; : alga ) is an informal term for any organisms of a large and diverse group of photosynthesis, photosynthetic organisms that are not plants, and includes species from multiple distinct clades. Such organisms range from unicellular ...
, to
protozoans Protozoa (: protozoan or protozoon; alternative plural: protozoans) are a polyphyletic group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic debris. Historically ...
, to
fungi A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
to
bacteria Bacteria (; : bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of Prokaryote, prokaryotic microorganisms. Typically a few micr ...
l or
microbial A microorganism, or microbe, is an organism of microscopic size, which may exist in its single-celled form or as a colony of cells. The possible existence of unseen microbial life was suspected from antiquity, with an early attestation in ...
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 Siphonophorae (from Ancient Greek σίφων (siphōn), meaning "tube" and -φόρος (-phóros), meaning "bearing") is an order within Hydrozoa, a class of marine organisms within the phylum Cnidaria. According to the World Register of Marine ...
, 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. On the early Earth, reactive elements, such as
iron Iron is a chemical element; it has symbol Fe () and atomic number 26. It is a metal that belongs to the first transition series and group 8 of the periodic table. It is, by mass, the most common element on Earth, forming much of Earth's o ...
and
uranium Uranium is a chemical element; it has chemical symbol, symbol U and atomic number 92. It is a silvery-grey metal in the actinide series of the periodic table. A uranium atom has 92 protons and 92 electrons, of which 6 are valence electrons. Ura ...
, existed in a reduced form that would react with any free oxygen produced by
photosynthesis Photosynthesis ( ) is a system of biological processes by which photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical energy necessary to fuel their metabo ...
ing organisms. Oxygen would not be able to build up in the
atmosphere An atmosphere () is a layer of gases that envelop an astronomical object, held in place by the gravity of the object. A planet retains an atmosphere when the gravity is great and the temperature of the atmosphere is low. A stellar atmosph ...
until all the iron had rusted (producing
banded iron formation Banded iron formations (BIFs; also called banded ironstone formations) are distinctive units of sedimentary rock consisting of alternating layers of iron oxides and iron-poor chert. They can be up to several hundred meters in thickness and e ...
s), 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 In physics, radiation is the emission or transmission of energy in the form of waves or particles through space or a material medium. This includes: * ''electromagnetic radiation'' consisting of photons, such as radio waves, microwaves, infr ...
. 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. However, if they were common, more than the occasional specimen might be expected in exceptionally preserved fossil assemblages (Konservat- Lagerstätten) such as the
Burgess Shale The Burgess Shale is a fossil-bearing deposit exposed in the Canadian Rockies of British Columbia, Canada. It is famous for the exceptional preservation of the soft parts of its fossils. At old (middle Cambrian), it is one of the earliest fos ...
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

It has been suggested that by the Early Cambrian, organisms higher in the
food chain A food chain is a linear network of links in a food web, often starting with an autotroph (such as grass or algae), also called a producer, and typically ending at an apex predator (such as grizzly bears or killer whales), detritivore (such as ...
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 Substrate may refer to: Physical layers *Substrate (biology), the natural environment in which an organism lives, or the surface or medium on which an organism grows or is attached ** Substrate (aquatic environment), the earthy material that exi ...
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 ''Kimberella'' is an extinct genus of marine bilaterian known only from rocks of the Ediacaran period. The slug-like organism fed by scratching the microbial surface on which it dwelt in a manner similar to the gastropods, although its affinity ...
'' as a grazer is correct then this suggests that the biota had already had limited exposure to "predation".


Competition

Increased competition due to the evolution of key innovations among other groups, perhaps as a response to predation, may have driven the Ediacaran biota from their niches. However, the supposed "competitive exclusion" of
brachiopod Brachiopods (), phylum (biology), phylum Brachiopoda, are a phylum of animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear e ...
s by
bivalve molluscs Bivalvia () or bivalves, in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of aquatic molluscs (marine and freshwater) that have laterally compressed soft bodies enclosed by a calcified exoskeleton consis ...
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
supercontinent In geology, a supercontinent is the assembly of most or all of Earth's continent, continental blocks or cratons to form a single large landmass. However, some geologists use a different definition, "a grouping of formerly dispersed continents", ...
s, 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 Marine chemistry, also known as ocean chemistry or chemical oceanography, is the study of the chemical composition and processes of the world’s oceans, including the interactions between seawater, the atmosphere, the seafloor, and marine organ ...
(promoting
biomineralisation Biomineralization, also written biomineralisation, is the process by which living organisms produce minerals, often resulting in hardened or stiffened '' mineralized tissues''. It is an extremely widespread phenomenon: all six taxonomic kingd ...
) 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.


Avalon assemblage

The Avalon assemblage is defined at Mistaken Point one the
Avalon Peninsula The Avalon Peninsula () is a large peninsula that makes up the southeast portion of the island of Newfoundland in Canada. It is in size. The peninsula is home to 270,348 people, about 52% of the province's population, according to the 2016 Ca ...
of Canada, the oldest locality with a large quantity of Ediacaran fossils. The assemblage is easily dated because it contains many fine ash-beds, which are a good source of
zircon Zircon () is a mineral belonging to the group of nesosilicates and is a source of the metal zirconium. Its chemical name is zirconium(IV) silicate, and its corresponding chemical formula is Zr SiO4. An empirical formula showing some of th ...
s used in the uranium-lead method of
radiometric dating Radiometric dating, radioactive dating or radioisotope dating is a technique which is used to Chronological dating, date materials such as Rock (geology), rocks or carbon, in which trace radioactive impurity, impurities were selectively incorporat ...
. 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
rangeomorph The rangeomorphs are a group of Ediacaran Ediacaran biota, fossils. Ediacarans are the oldest large fossil organisms on earth, and many are not self-evidently related to anything else that has ever lived. However, some Ediacarans clearly resemble ...
s such as '' Charnia'', all of which share a
fractal In mathematics, a fractal is a Shape, geometric shape containing detailed structure at arbitrarily small scales, usually having a fractal dimension strictly exceeding the topological dimension. Many fractals appear similar at various scale ...
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 The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the ...
suspension-feeding communities, which may suggest
filter feeding Filter feeders are aquatic animals that acquire nutrients by feeding on organic matters, food particles or smaller organisms (bacteria, microalgae and zooplanktons) suspended in water, typically by having the water pass over or through a spe ...
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 The White Sea (; Karelian language, Karelian and ; ) is a southern inlet of the Barents Sea located on the northwest coast of Russia. It is surrounded by Karelia to the west, the Kola Peninsula to the north, and the Kanin Peninsula to the nort ...
or Australia's
Ediacara Hills Ediacara Hills ( ), also known as Ediacaran Hills, are a range of low hills in the northern part of the Flinders Ranges of South Australia, around north of the state capital of Adelaide city centre, Adelaide. They are within the Nilpena Ediaca ...
and is marked by much higher diversity than the Avalon or Nama assemblages. In Australia, they are typically found in red gypsiferous and calcareous paleosols formed on
loess A loess (, ; from ) is a clastic rock, clastic, predominantly silt-sized sediment that is formed by the accumulation of wind-blown dust. Ten percent of Earth's land area is covered by loesses or similar deposition (geology), deposits. A loess ...
and flood deposits in an arid cool temperate paleoclimate. Most fossils are preserved as imprints in microbial beds, but a few are preserved ''within'' sandy units. (Source of data for Timeline synthesis, p. 218. Further citations available in caption to Fig. 8.)


Nama assemblage

The Nama assemblage is best represented in
Namibia Namibia, officially the Republic of Namibia, is a country on the west coast of Southern Africa. Its borders include the Atlantic Ocean to the west, Angola and Zambia to the north, Botswana to the east and South Africa to the south; in the no ...
. It is marked by extreme biotic turnover, with rates of extinction exceeding rates of origination for the whole period. Three-dimensional preservation is most common, with organisms preserved in sandy beds containing internal bedding. Dima Grazhdankin believes that these fossils represent burrowing organisms, while Guy Narbonne maintains they were surface dwellers. These beds are sandwiched between units comprising interbedded sandstones, siltstones and
shale Shale is a fine-grained, clastic sedimentary rock formed from mud that is a mix of flakes of Clay mineral, clay minerals (hydrous aluminium phyllosilicates, e.g., Kaolinite, kaolin, aluminium, Al2Silicon, Si2Oxygen, O5(hydroxide, OH)4) and tiny f ...
s—with microbial mats, where present, usually containing the fossils. The environment is interpreted as sand bars formed at the mouth of a
delta Delta commonly refers to: * Delta (letter) (Δ or δ), the fourth letter of the Greek alphabet * D (NATO phonetic alphabet: "Delta"), the fourth letter in the Latin alphabet * River delta, at a river mouth * Delta Air Lines, a major US carrier ...
's
distributaries A distributary, or a distributary channel is a stream channel that branches off and flows a main stream channel. It is the opposite of a ''tributary'', a stream that flows another stream or river. Distributaries are a result of river bifurca ...
. Mattress-like vendobionts ('' Ernietta'', '' Pteridinium'', '' Rangea'') in these sandstones form a very different assemblage from vermiform fossils ('' Cloudina'', ''
Namacalathus ''Namacalathus'' is a problematic metazoan fossil occurring in the latest Ediacaran. The first, and only described species, ''N. hermanastes'', was first described in 2000 from the Nama Group of central and southern Namibia. Discovery and nami ...
'') of Ediacaran "wormworld" in marine dolomite of Namibia.


Significance of assemblages

Since they are globally distributed – described on all continents except
Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean (also known as the Antarctic Ocean), it contains the geographic South Pole. ...
– 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 Continental drift is a highly supported scientific theory, originating in the early 20th century, that Earth's continents move or drift relative to each other over geologic time. The theory of continental drift has since been validated and inc ...
- an application of
paleomagnetism Paleomagnetism (occasionally palaeomagnetism) is the study of prehistoric Earth's magnetic fields recorded in rocks, sediment, or archeological materials. Geophysicists who specialize in paleomagnetism are called ''paleomagnetists.'' Certain ...
) and in separate
sedimentary basin Sedimentary basins are region-scale depressions of the Earth's crust where subsidence has occurred and a thick sequence of sediments have accumulated to form a large three-dimensional body of sedimentary rock They form when long-term subsidence ...
s. An analysis of one of the White Sea fossil beds, where the layers cycle from continental seabed to inter-tidal to estuarine and back again a few times, found that a specific set of Ediacaran organisms was associated with each environment. However, while there is some delineation in organisms adapted to different environments, the three assemblages are more distinct temporally than paleoenvironmentally. 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

* Cambrian explosion *
Large ornamented Ediacaran microfossil Large ornamented Ediacaran microfossils are microscopic acritarchs, usually over 100 μm in diameter, which are common in sediments of the Ediacaran period, . They largely disappear from the Ediacaran period fossil record before , roughly ...
*
List of Ediacaran genera The existence of life, especially that of animals, before the Cambrian had long been the subject of debate in paleontology. The apparent suddenness of the Cambrian explosion had no firm explanation, and Charles Darwin himself recognized the chal ...
*
Abiogenesis Abiogenesis is the natural process by which life arises from non-living matter, such as simple organic compounds. The prevailing scientific hypothesis is that the transition from non-living to living entities on Earth was not a single even ...
*
Huainan biota The Huainan biota is a collection of macroscopic organisms discovered in the early 1980s in the Precambrian deposits of China ( Huainan City, Anhui Province), that represent a transitional period in evolution. Discovery and Age Huainan biota org ...
* Francevillian biota, another, much earlier
Precambrian The Precambrian ( ; or pre-Cambrian, sometimes abbreviated pC, or Cryptozoic) is the earliest part of Earth's history, set before the current Phanerozoic Eon. The Precambrian is so named because it preceded the Cambrian, the first period of t ...
possibly pluricellular biota.


Notes


References


Further reading

* — Excellent further reading for the keen – includes many interesting chapters with macroevolutionary theme. * — A popular science account of these fossils, with a particular focus on the Namibian fossils. *


External links

* — Ediacaran biota review article, with nice illustrations * — Thorough, though slightly out-of-date, description * — compilation up to 7/2010 * * * * — ABC TV show on Ediacaran fossils at Nilpena * , 2009, radio program {{DEFAULTSORT:Ediacaran biota Biota Proterozoic life Precambrian paleobiotas