Ediacaran Biota

23/4/2020 Ediacaran biota - Wikipedia
Ediacaran biota
The Ediacaran (/ˌiːdiˈækərən/; formerly Vendian) biota
is a taxonomic period classification that consists of all life
forms that were present on Earth during the Ediacaran
Period (ca. 635–542 Mya). These were composed of
enigmatic tubular and frond-shaped, mostly sessile,
organisms. Trace fossils of these organisms have been found
worldwide, and represent the earliest known complex
multicellular organisms.[note 1]
The Ediacaran biota may have undergone evolutionary

radiation in a proposed event called the Avalon explosion,
575 million years ago.[1][2] This was after the Earth had
thawed from the Cryogenian period's extensive glaciation. Dickinsonia costata, an Ediacaran organism,
This biota largely disappeared with the rapid increase in displays the characteristic quilted appearance
biodiversity known as the Cambrian explosion. Most of the of Ediacaran enigmata
currently existing body plans of animals first appeared in
the fossil record of the Cambrian rather than the Ediacaran.
For macroorganisms, the Cambrian biota appears to have 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 600 million years ago and flourished until
the cusp of the Cambrian 542 million years ago, when the characteristic communities of fossils vanished.
A diverse Ediacaran community was discovered in 1995 in Sonora, Mexico, and is approximately 555
million years in age, roughly coeval with Ediacaran fossils of the Ediacara Hills, South Australia and the
White Sea, Russia.[3][4][5] While rare fossils that may represent survivors have been found as late as the
Middle Cambrian (510 to 500 million years ago), the earlier fossil communities disappear from the
record at the end of the Ediacaran leaving only curious fragments of once-thriving ecosystems.[6]
Multiple hypotheses exist to explain the disappearance of this biota, including preservation bias, a
changing environment, the advent of predators and competition from other life-forms. Recent (2018)
sampling of late Ediacaran strata across Baltica (<560 mya) 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.[7]
Determining where Ediacaran organisms fit in the tree of life has proven challenging; it is not even
established that they were animals, with suggestions that they were lichens (fungus-alga symbionts),
algae, protists known as foraminifera, fungi or microbial colonies, or hypothetical intermediates between
plants and animals.[8] The morphology and habit of some taxa (e.g. Funisia dorothea) suggest
relationships to Porifera or Cnidaria.[9] Kimberella may show a similarity to molluscs, and other
organisms have been thought to possess bilateral symmetry, 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 palaeontologists have suggested that these represent completely extinct lineages
that do not resemble any living organism. One palaeontologist proposed a separate kingdom level
category Vendozoa (now renamed Vendobionta)[10] in the Linnaean hierarchy for the Ediacaran
biota. If these enigmatic organisms left no descendants, their strange forms might be seen as a "failed
https://en.wikipedia.org/wiki/Ediacaran_biota 1/26
experiment" in multicellular life, with later multicellular life evolving independently from unrelated
single-celled organisms.[11] However, a 2018 study confirmed that one of the period's most-prominent
and iconic fossils, Dickinsonia, included cholesterol,[12] limiting its affinities to that of animals, fungi, or
red algae.[13]
The concept of "Ediacaran Biota" is somewhat

artificial as it can not be defined geographically, The Ediacara biota in context
stratigraphically, taphonomically, or biologically.[16]
-490 —
–
-500 — ←Last putative
Ediacaran
Contents –
-510 —
History – Cambrian
-520 —
Preservation –
Microbial mats -530 —
Fossilization –
-540 —
Scale of preservation ←Last Ediacaran
– communities
Morphology -550 —
–
Classification and interpretation Charnia
-560 —
Cnidarians –
Protozoans -570 —
New phylum – ←Avalon explosion
-580 —
Lichen hypothesis ←Gaskiers
– Aspidella
Other interpretations discs Glaciation
-590 —
Origin –
-600 —
Preservation bias
–
Predation and grazing -610 — ←First Ediacaran
Competition – megafossil
-620 — Ediacaran
Change in environmental conditions
–
Assemblages -630 —
←Embryos?
Avalon-type assemblage – ←Last extensive
Ediacara-type assemblage -640 — glaciation
– Cryogenian
Nama-type assemblage -650 —
Significance of assemblages
Neoproterozoic
See also
Notes (last era of the Precambrian)
Palaeozoic
References (first era of the Phanerozoic)
Further reading
Axis scale: million years
External links
References: Waggoner 1998,[14] Hofmann
1990[15]
History
Life timeline
The first Ediacaran fossils discovered were the Ice Ages

0—
Quaternary Primates ←Earliest apes
disc-shaped Aspidella terranovica in 1868. Their P
Flowers
Birds Mammals
h
discoverer, Scottish geologist Alexander Murray, Karoo – a Plants Dinosaurs
found them useful aids for correlating the age of Andean n ←Tetrapoda
e
rocks around Newfoundland.[17] However, since -500 — r
Arthropods Molluscs
←Cambrian explosion
o ←Ediacara biota
they lay below the "Primordial Strata" of the Cryogenian
– z ←Earliest animals
o
Cambrian that was then thought to contain the i ←Earliest plants
very first signs of animal life, a proposal four -1000 — c Multicellular
years after their discovery by Elkanah Billings life
–
←Sexual reproduction
that these simple forms represented fauna was P
dismissed by his peers. Instead, they were -1500 —
r
o
interpreted as gas escape structures or inorganic t
concretions.[17] No similar structures elsewhere – e
r Eukaryotes
in the world were then known and the one-sided o
-2000 — z
debate soon fell into obscurity.[17] In 1933, Georg o
i
Gürich discovered specimens in Namibia[18] but Huronian – c
←Oxygen crisis
the firm belief that complex life originated in the
←Atmospheric oxygen
Cambrian led to them being assigned to the -2500 —
Cambrian Period and no link to Aspidella was Photosynthesis
–
made. In 1946, Reg Sprigg noticed "jellyfishes" in Pongola
the Ediacara Hills of Australia's Flinders -3000 — A

Ranges[19] but these rocks were believed to be r
c
Early Cambrian so, while the discovery sparked – h
e
some interest, little serious attention was a
-3500 — n ←Earliest oxygen
garnered.
Single-celled
– life
-4000 — ←Earliest life
H
a water
–d
e
a ←Earliest water
-4500 — n ←Earth (−4540)
(million years ago)
(
Palaeontologist Guy Narbonne
examining Ediacaran fossils in It was not until the British discovery of the iconic Charnia that the
Newfoundland pre-Cambrian 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 (who was not believed) and then the
next year by a group of three schoolboys including 15-year-old Roger Mason[20][21][22] and 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[23][24] and with a combination of improved dating of existing specimens and an
injection of vigour into the search many more instances were recognised.[25]
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.[26][27] It was also the first
discovery of Ediacarans in deep water sediments.[28]
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"[29] 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.[30]
The term "Ediacaran biota" and similar ("Ediacara"/"Ediacaran"/"Ediacarian"/"Vendian",

"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.[31]
Preservation
Microbial mats
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.[32]
Modern cyanobacterial-algal mat,
salty lake on the White Sea seaside
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, the evolution of grazing organisms in the Cambrian vastly reduced their
numbers.[33] 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.[34]
Fossilization
The preservation of these fossils is one of their great fascinations to science. As soft-bodied organisms,
they would normally not fossilize and, 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 were a global phenomenon. The processes that were operating must have been
systemic and worldwide. There was something very different about the Ediacaran Period that permitted
these delicate creatures to be left behind and it is thought the fossils were preserved by virtue of rapid
covering by ash or sand, trapping them against the mud or microbial mats on which they lived.[35] 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.[36] Ash beds provide more detail
and can readily be dated to the nearest million years or better using radiometric dating.[37] However, it is
more common to find Ediacaran fossils under sandy beds deposited by storms or high-energy bottom-
scraping ocean currents known as turbidites.[35] 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.[38]
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 The fossil Charniodiscus is
not destroy them as it would have the less-resistant discs. Further, in some barely distinguishable from
cases, the bacterial precipitation of minerals formed a "death mask", the "elephant skin" texture
on this cast.
ultimately leaving a positive, cast-like impression of the organism.[39][40]
Morphology
The Ediacaran biota exhibited a vast range of Forms of Ediacaran fossil
morphological characteristics. Size ranged from The earliest
millimetres to metres; complexity from "blob-like" to
discovered
intricate; rigidity from sturdy and resistant to jelly-soft.
potential
Almost all forms of symmetry were present.[35] These
organisms differed from earlier fossils by displaying an embryo,
organised, differentiated multicellular construction and preserved
centimetre-plus sizes. within an
acanthomorphic
These disparate morphologies can be broadly grouped acritarch. The
into form taxa: term 'acritarch'
"Embryos" describes a
Recent discoveries of Precambrian multicellular life range of
have been dominated by reports of embryos, unclassified
particularly from the Doushantuo Formation in cell-like fossils.
China. Some finds[41] generated intense media Tateana inflata
excitement[42] though some have claimed they are
(=
instead inorganic structures formed by the
'Cyclomedusa'
precipitation of minerals on the inside of a hole.[43]
Other "embryos" have been interpreted as the radiata) is the
remains of the giant sulfur-reducing bacteria akin attachment disk
to Thiomargarita,[44] a view that, while it had of an unknown
enjoyed a notable gain of supporters[45][46] as of organism.
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 A cast of the

decay.[47] quilted Charnia,
Microfossils dating from 632.5 million years ago – the first
just 3 million years after the end of the Cryogenian accepted
glaciations – may represent embryonic 'resting
stages' in the life cycle of the earliest known complex
animals.[48] An alternative proposal is that these Precambrian
structures represent adult stages of the organism.
multicellular organisms of this period.[49] Charnia was
once
Discs interpreted as a
Circular fossils, such as Ediacaria, Cyclomedusa
relative of the
and Rugoconites led to the initial identification of
Ediacaran fossils as cnidaria, which include sea pens.
jellyfish and corals.[19] Further examination has Spriggina was
provided alternative interpretations of all disc- originally
shaped fossils: not one is now confidently interpreted as
recognised as a jellyfish. Alternate explanations
annelid or
include holdfasts and protists;[50] the patterns
arthropod.
displayed where two meet have led to many
'individuals' being identified as microbial However, lack
colonies,[51][52] and yet others may represent of known limbs,
scratch marks formed as stalked organisms spun and glide
around their holdfasts.[53] Useful diagnostic reflected
characters are often lacking because only the isomers instead
underside of the organism is preserved by of true
fossilisation.
segments,
Bags rejects any
Fossils such as Pteridinium preserved within such
sediment layers resemble "mud-filled bags". The classification
scientific community is a long way from reaching a despite some
consensus on their interpretation.[54] superficial
Toroids resemblance.
The fossil Vendoglossa tuberculata from the Nama Late Ediacaran
Group, Namibia, has been interpreted as a dorso- Archaeonassa-
ventrally compressed stem-group metazoan, with a type trace
large gut cavity and a transversely ridged
fossils are
ectoderm. The organism is in the shape of a
flattened torus, with the long axis of its toroidal commonly
body running through the approximate center of preserved on
the presumed gut cavity.[55] the top surfaces
of sandstone
Quilted organisms strata.
The organisms considered in Seilacher's revised
definition of the Vendobionta[10] share a "quilted"
appearance and resembled an inflatable mattress. Epibaion
Sometimes these quilts would be torn or ruptured waggoneris,
prior to preservation: such damaged specimens chain of trace
provide valuable clues in the reconstruction platforms and
process. For example, the three (or more) petaloid
the imprint of
fronds of Swartpuntia germsi could only be
recognised in a posthumously damaged the body of

specimen – usually multiple fronds were hidden as Yorgia
burial squashed the organisms flat.[56] waggoneri
(right), which
These organisms appear to form two groups: the
fractal rangeomorphs and the simpler created these
erniettomorphs.[57] Including such fossils as the traces on
iconic Charnia and Swartpuntia, the group is both microbial mat.
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[58] or osmosis.[59] However, others argue against this.[60]
Non-Vendobionts
Some Ediacaran organisms have more complex details preserved, which has allowed them to be
interpreted as 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.[43][61] Later examples are almost universally
accepted as bilaterians and include the mollusc-like Kimberella,[62] Spriggina (pictured)[63] and the
shield-shaped Parvancorina[64] whose affinities are currently debated.[65]
A suite of fossils known as the Small shelly fossils are represented in the Ediacaran, most
famously by Cloudina[66] 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.
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.[67]
Fossils of the hard-shelled foraminifera Platysolenites are known from the latest Ediacaran of
western Siberia, coexisting with Cloudina and Namacalathus.[68]
Trace fossils
With the exception of some very simple vertical burrows[69] 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[70] but they could also have been
made by simpler organisms feeding as they slowly rolled along the sea floor.[71] Putative "burrows"
dating as far back as 1,100 million years may have been made by animals that fed on the
undersides of microbial mats, which would have shielded them from a chemically unpleasant
ocean;[72] however their uneven width and tapering ends make a biological origin so difficult to
defend[73] that even the original proponent no longer believes they are authentic.[74]
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;[75] 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.[76]
The potential mollusc Kimberella is associated with scratch marks, perhaps formed by a radula.[77]
Classification and interpretation

Classification of the Ediacarans is difficult, and 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.[78]
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".[50]
In 2018 analysis of ancient sterols was taken as evidence that one of the period's most-prominent and
iconic fossils, Dickinsonia, was an early animal.[12]
Cnidarians
Since the most primitive eumetazoans—multi-cellular animals with

tissues—are cnidarians, the first attempt to categorise these fossils
designated them as jellyfish and sea pens.[79] 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.[80][81]
A sea pen, a modern cnidarian
The link between certain frond-like Ediacarans and sea pens has
bearing a passing resemblance to
been thrown into doubt by multiple lines of evidence; chiefly the
Charnia
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.[82] Some researchers have suggested that an analysis of "growth poles" discredits the
pennatulacean nature of Ediacaran fronds.[83][84]
Protozoans
Adolf Seilacher has suggested the Ediacaran sees animals usurping giant protists as the dominant life
form.[85] The modern xenophyophores are giant single-celled protozoans found throughout the world's
oceans, largely on the abyssal plain. A recent genetic study suggested that the xenophyophores are a
specialised group of Foraminifera. There are approximately 42 recognised species in 13 genera and 2
orders; one of which, Syringammina fragilissima, is among the largest known protozoans at up to 20
centimetres in diameter.
New phylum
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,[86][87]
named after the now-obsolete Vendian era. He later excluded fossils
identified as metazoans and relaunched the phylum "Vendobionta".
He described the Vendobionta as quilted cnidarians lacking stinging

cells. This absence precludes the current cnidarian method of
feeding, so Seilacher suggested that the organisms may have
A single-celled xenophyophore in
survived by symbiosis with photosynthetic or chemoautotrophic
the Galapagos Rift
organisms.[88] 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".[89]
Lichen hypothesis
Greg Retallack's hypothesis that

Ediacaran organisms were
lichens [91][92] has been
controversial.[93][94][95] He argues that
the fossils are not as squashed as
known fossil jellyfish, and their relief is
closer to compressed woody branches
whose compaction can be estimated as
compressed cylinders. He points out Thin sections and substrates of a
the chitinous walls of lichen colonies variety of Ediacaran fossils [90]
would provide a similar resistance to
compaction, and claims the large size of
A modern lichen, the organisms (up to 1.5 metres long, far larger than any of the preserved
Hypogymnia burrows) also hints against classification with animals. Thin sections of
Ediacaran fossils show lichen-like compartments and hypha-like wisps of
ferruginized clay.[90] Finally, Ediacaran fossils from classic localities of the
Flinders Ranges have been found in strata that Rettalack interprets to be both growth position within
strata that he controversially interprets to be red calcareous and gypsiferous paleosols and possibly well-
drained temperate desert soils.[92][96] According to Retallack's interpretations, such habitats limit
interpretive options for fractal Ediacaran fossils such as Dickinsonia to lichenised or unlichenised fungi,
but other Ediacaran fossils could have been slime moulds or microbial colonies.
Other interpretations
Several classifications have been used to accommodate the Ediacaran biota at some point,[97] from
algae,[98] to protozoans,[99] to fungi[100] to bacterial or microbial colonies,[51] to hypothetical
intermediates between plants and animals.[8]
A new extant genus discovered in 2014, Dendrogramma, which appears to be a basal metazoan but of
unknown taxonomic placement, has been noted to have similarities with the Ediacaran fauna.[101] It has
since been found to be a siphonophore, possibly even sections of a more complex species,[102] though
this in turn has raised suspicions for a similar status for at least some ediacaran organisms.
Origin
It took almost 4 billion years from the formation of the Earth for the Ediacaran fossils to first appear,
655 million years ago. While putative fossils are reported from 3,460 million years ago,[103][104] the first
uncontroversial evidence for life is found 2,700 million years ago,[105] and cells with nuclei certainly
existed by 1,200 million years ago:[106] The reason why it took so long for forms with an Ediacaran grade
of organisation to appear is uncertain.
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
610 million year old Twitya formation,[107], and older rocks dating to 770 million years ago in
Kazakhstan [108], possibly displaying the most complex morphology of the time.
The alternative train of thought is that it was simply not

advantageous to be large until the appearance of the Ediacarans: the
environment favoured the small over the large. Examples of such
scenarios today include plankton, whose small size allows them to
reproduce rapidly to take advantage of ephemerally abundant
nutrients in algal blooms. But for large size never to be favourable,
the environment would have to be very different indeed.
A primary size-limiting factor is the amount of atmospheric oxygen.

Without a complex circulatory system, low concentrations of oxygen Global ice sheets may have delayed
cannot reach the centre of an organism quickly enough to supply its or prevented the establishment of
metabolic demand. 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[109] – and the presence of atmospheric oxygen was soon
heralded as a possible trigger for the Ediacaran radiation.[110] 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.[111]
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.[112]
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.[113] Potentially, complex life may have evolved before these glaciations,
and been wiped out. However, the diversity of life in modern Antarctica has sparked disagreement over
whether cold temperatures increase or decrease the rate of evolution.
In early 2008 a team analysed the range of basic body structures ("disparity") of Ediacaran organisms
from three different fossil beds: Avalon in Canada, 575 to 565 million years ago; White Sea in Russia,
560 to 550 million years ago; and Nama in Namibia, 550 to 542 million years ago, 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 .[114]
Preservation bias
The paucity of Ediacaran fossils after the Cambrian could simply be due to conditions that no longer
favoured the fossilisation of Ediacaran organisms, which may have continued to thrive unpreserved.[32]
However, if they were common, more than the occasional specimen[6][115] might be expected in
exceptionally preserved fossil assemblages (Konservat-Lagerstätten) such as the Burgess Shale and
Chengjiang.[116] There are at present no widely accepted reports of Ediacara-type organisms in the
Cambrian period, though there are a few disputed reports, as well as unpublished observations of
'vendobiont' fossils from 535 Ma Orsten-type deposits in China.[117]
Predation and grazing
It is 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 substrate, leading
to displacement or detachment of the biota; or that the destruction
of the mat destabilised the ecosystem, causing extinctions.
Alternatively, skeletonised animals could have fed directly on the

relatively undefended Ediacaran biota.[50] 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 Kimberella may have had a
"predation".[62] predatory or grazing lifestyle.
There is however little evidence for any trace fossils in the Ediacaran
Period, which may speak against the active grazing theory. Further, the onset of the Cambrian Period is
defined by the appearance of a worldwide trace fossil assemblage, quite distinct from the activity-barren
Ediacaran Period.
Competition
It is possible that increased competition due to the evolution of key

innovations among other groups, perhaps as a response to
predation,[118] drove the Ediacaran biota from their niches.
However, this argument has not successfully explained similar
phenomena. For instance, the bivalve molluscs' "competitive
exclusion" of brachiopods was eventually deemed to be a
coincidental result of two unrelated trends.[119]
Cambrian animals such as Waptia
may have competed with, or fed
Change in environmental conditions
upon, Ediacaran life-forms.
While it is difficult to infer the effect of changing planetary conditions on organisms, communities and
ecosystems, great changes were happening at the end of the Precambrian and the start of the Early
Cambrian. The breakup of the supercontinents,[120] rising sea levels (creating shallow, "life-friendly"
seas),[121] a nutrient crisis,[122] fluctuations in atmospheric composition, including oxygen and carbon
dioxide levels,[123] and changes in ocean chemistry[124] (promoting biomineralisation)[125] could all have
played a part.
Assemblages
Ediacaran-type fossils are recognised globally in 25 localities[30] and a variety of depositional conditions,
and are commonly grouped into three main types, known as assemblages and named after typical
localities. Each assemblage tends to occupy its own region of morphospace, and after an initial burst of
diversification changes little for the rest of its existence.[126]
Avalon-type assemblage
The Avalon-type assemblage is defined at Mistaken Point in Canada,

the oldest locality with a large quantity of Ediacaran fossils.[128] 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.[126]
One interpretation of the biota is as deep-sea-dwelling

rangeomorphs[129] 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 Ediacara- or
Nama-types, resembles Carboniferous suspension-feeding
communities, which may suggest filter feeding [130] – by most
Reconstruction of fossil soils and
their biota in the Mistaken Point interpretations, the assemblage is found in water too deep for
Formation of Newfoundland [127] photosynthesis. The low diversity may reflect the depth of water –
which would restrict speciation opportunities – or it may just be too
young for a rich biota to have evolved. Opinion is currently divided
between these conflicting hypotheses.[131]
An alternative explanation for the distinct composition of the Avalon-type assemblage is that it was a
terrestrial assemblage of volcaniclastic coastal soils near a continental volcanic arc.[127] This view is
based on geochemical studies of the substrates of Mistaken Point fossils and associated matrix supported
tuffs and volcanic bombs that could only form on land.[132] Some of these fossils such as Fractofusus and
Charniodiscus were found in strata that Retallack interprets to be red well drained paleosols of coastal
plains, but others such as Aspidella were found in strata that Retallack interprets to be intertidal
paleosols.
Ediacara-type assemblage
The Ediacara-type assemblage is named after Australia's Ediacara

Hills, and consists of fossils preserved in facies of coastal lagoons
and rivers. They are typically found in red gypsiferous and
calcareous paleosols formed on loess and flood deposits in an arid
cool temperate paleoclimate.[92] Most fossils are preserved as
imprints in microbial earths,[133] but a few are preserved within
sandy units.[131][126]
Biota ranges[131]
Reconstruction of Ediacaran biota

and their soils in the Ediacara
Member of the Rawnsley Quartzite
in the Flinders Ranges, South
Australia [92]
Nama-type assemblage
The Nama assemblage is best represented

in Namibia. Three-dimensional
preservation is most common, with
organisms preserved in sandy beds
containing internal bedding. Dima
Grazhdankin believes that these fossils
represent burrowing organisms,[54] while
Guy Narbonne maintains they were surface
dwellers.[134] These beds are sandwiched
between units comprising interbedded
sandstones, siltstones and shales – with Axis scale: millions of years ago, dated with U/Pb of zircons
microbial mats, where present, usually
containing the fossils. The environment is interpreted as sand bars formed at the mouth of a delta's
distributaries.[131] Mattress-like vendobionts (Ernietta, Pteridinium, Rangea) in these sandstones form
a very different assemblage from vermiform fossils (Cloudina, Namacalathus) of Ediacaran
"wormworld" in marine dolomite of Namibia.[135]
Significance of assemblages
In the White Sea region of Russia, all three assemblage types have been found in close proximity. This,
and the faunas' considerable temporal overlap, makes it unlikely that they represent evolutionary stages
or temporally distinct communities. Since they are globally distributed – described on all continents
except Antarctica – geographical boundaries do not appear to be a factor;[136] the same fossils are found
at all palaeolatitudes (the latitude where the fossil was created, accounting for continental drift) and in
separate sedimentary basins.[131]
It is most likely that the three assemblages mark organisms adapted to survival in different
environments, and that any apparent patterns in diversity or age are in fact an artefact of the few
samples that have been discovered – the timeline (right) demonstrates the paucity of Ediacaran fossil-
bearing assemblages. 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.[131]
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.[137] The lack of large-scale
predation and vertical burrowing are perhaps the most significant factors limiting the ecological
diversity; the emergence of these during the Early Cambrian allowed the number of lifestyles occupied to
rise to 30.
See also
Cambrian explosion
Large ornamented Ediacaran microfossil
List of Ediacaran genera
Origin of life
Francevillian biota
Notes
1. Simple multicellular organisms such as red algae evolved at least 1,200 million years ago. The
status of the Francevillian biota of 2,100 million years ago is unclear, but they may represent earlier
multicellular forms of a more complex nature.
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1a7be). GSA Today. 26 (11): 4–11. doi:10.1130/GSATG265A.1 (https://doi.org/10.1130%2FGSATG2
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org/10.1111%2Fj.1475-4983.2006.00611.x).
Further reading
Derek Briggs; Peter Crowther, eds. (2001). Palæobiology II: A synthesis. Malden, MA: Blackwell
Science. pp. Chapter 1. ISBN 978-0-632-05147-2. OCLC 43945263 (https://www.worldcat.org/oclc/4
3945263). Excellent further reading for the keen – includes many interesting chapters with
macroevolutionary theme.
Mark McMenamin (1998). The Garden of Ediacara: Discovering the First Complex Life. New York:
Columbia University Press. pp. 368pp. ISBN 978-0-231-10558-3. OCLC 3758852 (https://www.world
cat.org/oclc/3758852). A popular science account of these fossils, with a particular focus on the
Namibian fossils.
Wood, Rachel A., "The Rise of Animals: New fossils and analyses of ancient ocean chemistry reveal
the surprisingly deep roots of the Cambrian explosion", Scientific American, vol. 320, no. 6 (June
2019), pp. 24–31.
External links
Ediacara Biota (https://www.bbc.co.uk/programmes/b00lh2s3) on In Our Time at the BBC
"The oldest complex animal fossils" (https://web.archive.org/web/20070528175308/http://geol.queen
su.ca/museum/exhibits/ediac/drook/) – Queen's University, Canada
"Ediacaran fossils of Canada" (https://web.archive.org/web/20070405091608/http://geol.queensu.ca/

museum/exhibits/ediac/ediac.html) – Queen's University, Canada
"The Ediacaran Assemblage" (https://web.archive.org/web/20101010035019/http://www.peripatus.ge
n.nz/paleontology/Ediacara.html) – Thorough, though slightly out-of-date, description
"Database of Ediacaran Biota" (https://web.archive.org/web/20110725191154/http://www.complex-lif
e.org/database) Advent of Complex Life
Earth's oldest animal ecosystem held in fossils at Nilpena Station in SA outback (http://www.abc.net.
au/news/2013-08-03/sa-pastoral-property-nilpena-holds-animal-fossils/4862432) ABC News, 5
August 2013. Accessed 6 August 2013.
Meet the fossils (http://www.abc.net.au/tv/programs/landline/old-site/content/2013/s3817544.htm)
ABC Landline TV program on Ediacaran fossils at Nilpena (audio + transcript). First broadcast 3
August 2013. Accessed 28 December 2018.
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23/4/2020 Ediacaran biota - Wikipedia

The Ediacaran biota may have undergone evolutionary

The concept of "Ediacaran Biota" is somewhat

The first Ediacaran fossils discovered were the Ice Ages

the Ediacara Hills of Australia's Flinders -3000 — A

The term "Ediacaran biota" and similar ("Ediacara"/"Ediacaran"/"Ediacarian"/"Vendian",

Microbial mats are areas of sediment stabilised by the presence of

scraping ocean currents known as turbidites.[35] Soft-bodied organisms

The rate of cementation of the overlying substrate relative to the rate of

Conversely, quilted fossils tended to decompose after the cementation of

specimens, both living and in various stages of A cast of the

recognised in a posthumously damaged the body of

Classification and interpretation

Since the most primitive eumetazoans—multi-cellular animals with

Seilacher has suggested that the Ediacaran organisms represented a

He described the Vendobionta as quilted cnidarians lacking stinging

Greg Retallack's hypothesis that

The alternative train of thought is that it was simply not

A primary size-limiting factor is the amount of atmospheric oxygen.

On the early Earth, reactive elements, such as iron and uranium,

Predation and grazing

It is suggested that by the Early Cambrian, organisms higher in the

Alternatively, skeletonised animals could have fed directly on the

It is possible that increased competition due to the evolution of key

The Avalon-type assemblage is defined at Mistaken Point in Canada,

One interpretation of the biota is as deep-sea-dwelling

The Ediacara-type assemblage is named after Australia's Ediacara

Reconstruction of Ediacaran biota

The Nama assemblage is best represented

"Ediacaran fossils of Canada" (https://web.archive.org/web/20070405091608/http://geol.queensu.ca/

Retrieved from "https://en.wikipedia.org/w/index.php?title=Ediacaran_biota&oldid=952589057"

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