Cambrian explosion

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Not to be confused with Cambrian Colliery.

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The Cambrian explosion

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The Cambrian explosion or Cambrian radiation[1] was an event

approximately 541 million years ago in the Cambrian period when practically all
major animal phyla started appearing in the fossil record.[2][3] It lasted for about 13[4][5]
[6]
 – 25[7][8] million years and resulted in the divergence of most
modern metazoan phyla.[9] The event was accompanied by major diversification of
other organisms.[a]
Before the Cambrian explosion,[b] most organisms were simple, composed of
individual cells occasionally organized into colonies. As the rate of diversification
subsequently accelerated, the variety of life began to resemble that of today.
[11]
 Almost all present animal phyla appeared during this period. [12][13]
Key Cambrian explosion events
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Ediacaran
Biota
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Baykonur
glaciation
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Orsten Fauna
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Burgess Shale
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Kaili biota
←
Archaeocyatha extinction
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Emu Bay Shale
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Sirius Passet biota
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Chengjiang biota
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First arthropods with mineralized carapace (Trilobites)
←
SSF diversification, first brachiopods & archaeocyatha
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First halkieriids, mollusсs, hyoliths SSF
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Treptichnus pedum
Large negative δ13C peak
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First Cloudina & Namacalathus mineral tubular fossils
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Mollusc-like Kimberella and its trace fossils
←
Gaskiers glaciation
Archaeonassa-type trace fossils

Contents

 1History and significance

o 1.1Dating the Cambrian
o 1.2Body fossils
o 1.3Trace fossils
o 1.4Geochemical observations
o 1.5Phylogenetic techniques
 2Explanation of key scientific terms
o 2.1Phylum
o 2.2Stem group
o 2.3Triploblastic
o 2.4Bilaterian
o 2.5Coelomate
 3Precambrian life
o 3.1Evidence of animals around 1 billion years ago
o 3.2Fossils of the Doushantuo formation
o 3.3Burrows
o 3.4Ediacaran organisms
o 3.5Beck Spring Dolomite
 4Ediacaran–Early Cambrian skeletonisation
 5Cambrian life
o 5.1Trace fossils
o 5.2Small shelly fauna
o 5.3Early Cambrian trilobites and echinoderms
o 5.4Burgess Shale type faunas
o 5.5Early Cambrian crustaceans
o 5.6Early Ordovician radiation
 6Stages
 7Validity
o 7.1The Cambrian explosion as survivorship bias
 8Possible causes
o 8.1Changes in the environment
  8.1.1Increase in oxygen levels
 8.1.2Ozone formation
 8.1.3Snowball Earth
 8.1.4Increase in the calcium concentration of the Cambrian
seawater
o 8.2Developmental explanations
o 8.3Ecological explanations
 8.3.1End-Ediacaran mass extinction
 8.3.2Anoxia
 8.3.3Evolution of eyes
 8.3.4Arms races between predators and prey
 8.3.5Increase in size and diversity of planktonic animals
o 8.4Ecosystem engineering
o 8.5Complexity threshold
 9Uniqueness of the explosion
 10See also
 11Footnotes
 12References
 13Further reading
 14External links

History and significance[edit]

Life timeline
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(See also: Human timeline, and Nature timeline.)

Main article: Evolutionary history of life

The seemingly rapid appearance of fossils in the "Primordial Strata" was noted
by William Buckland in the 1840s,[14] and in his 1859 book On the Origin of
Species, Charles Darwin discussed the then inexplicable lack of earlier fossils as
one of the main difficulties for his theory of descent with slow modification
through natural selection.[15] The long-running puzzlement about the appearance of
the Cambrian fauna, seemingly abruptly, without precursor, centers on three key
points: whether there really was a mass diversification of complex organisms over a
relatively short period of time during the early Cambrian; what might have caused
such rapid change; and what it would imply about the origin of animal life.
Interpretation is difficult due to a limited supply of evidence, based mainly on an
incomplete fossil record and chemical signatures remaining in Cambrian rocks.
The first discovered Cambrian fossils were trilobites, described by Edward Lhuyd,
the curator of Oxford Museum, in 1698.[16] Although their evolutionary importance was
not known, on the basis of their old age, William Buckland (1784–1856) realized that
a dramatic step-change in the fossil record had occurred around the base of what we
now call the Cambrian.[14] Nineteenth-century geologists such as Adam
Sedgwick and Roderick Murchison used the fossils for dating rock strata, specifically
for establishing the Cambrian and Silurian periods.[17] By 1859, leading geologists
including Roderick Murchison, were convinced that what was then called the lowest
Silurian stratum showed the origin of life on Earth, though others, including Charles
Lyell, differed. In On the Origin of Species, Charles Darwin considered this sudden
appearance of a solitary group of trilobites, with no apparent antecedents, and
absence of other fossils, to be "undoubtedly of the gravest nature" among the
difficulties in his theory of natural selection. He reasoned that earlier seas had
swarmed with living creatures, but that their fossils had not been found due to the
imperfections of the fossil record.[15] In the sixth edition of his book, he stressed his
problem further as:[18]
To the question why we do not find rich fossiliferous deposits belonging to these
assumed earliest periods prior to the Cambrian system, I can give no satisfactory
answer.
American paleontologist Charles Walcott, who studied the Burgess Shale fauna,
proposed that an interval of time, the "Lipalian", was not represented in the fossil
record or did not preserve fossils, and that the ancestors of the Cambrian animals
evolved during this time.[19]
Earlier fossil evidence has since been found. The earliest claim is that the history of
life on earth goes back 3,850 million years:[20] Rocks of that age at Warrawoona,
Australia, were claimed to contain fossil stromatolites, stubby pillars formed by
colonies of microorganisms. Fossils (Grypania) of more complex eukaryotic cells,
from which all animals, plants, and fungi are built, have been found in rocks
from 1,400 million years ago, in China and Montana. Rocks dating
from 580 to 543 million years ago contain fossils of the Ediacara biota, organisms so
large that they are likely multicelled, but very unlike any modern organism. [21] In
1948, Preston Cloud argued that a period of "eruptive" evolution occurred in the
Early Cambrian,[22] but as recently as the 1970s, no sign was seen of how the
'relatively' modern-looking organisms of the Middle and Late Cambrian arose.[21]

Opabinia made the largest single contribution to modern interest in the Cambrian explosion.

The intense modern interest in this "Cambrian explosion" was sparked by the work
of Harry B. Whittington and colleagues, who, in the 1970s, reanalysed many fossils
from the Burgess Shale and concluded that several were as complex as, but
different from, any living animals.[23][24] The most common organism, Marrella, was
clearly an arthropod, but not a member of any known arthropod class. Organisms
such as the five-eyed Opabinia and spiny slug-like Wiwaxia were so different from
anything else known that Whittington's team assumed they must represent different
phyla, seemingly unrelated to anything known today. Stephen Jay Gould's popular
1989 account of this work, Wonderful Life,[25] brought the matter into the public eye
and raised questions about what the explosion represented. While differing
significantly in details, both Whittington and Gould proposed that all modern animal
phyla had appeared almost simultaneously in a rather short span of geological
period. This view led to the modernization of Darwin's tree of life and the theory
of punctuated equilibrium, which Eldredge and Gould developed in the early 1970s
and which views evolution as long intervals of near-stasis "punctuated" by short
periods of rapid change.[26]
Other analyses, some more recent and some dating back to the 1970s, argue that
complex animals similar to modern types evolved well before the start of the
Cambrian.[27][28][29]
Dating the Cambrian[edit]
Radiometric dates for much of the Cambrian, obtained by analysis of radioactive
elements contained within rocks, have only recently become available, and for only a
few regions.
Relative dating (A was before B) is often assumed sufficient for studying processes
of evolution, but this, too, has been difficult, because of the problems involved in
matching up rocks of the same age across different continents.[30]
Therefore, dates or descriptions of sequences of events should be regarded with
some caution until better data becomes available.
Body fossils[edit]
Fossils of organisms' bodies are usually the most informative type of evidence.
Fossilization is a rare event, and most fossils are destroyed
by erosion or metamorphism before they can be observed. Hence, the fossil record
is very incomplete, increasingly so as earlier times are considered. Despite this, they
are often adequate to illustrate the broader patterns of life's history. [31] Also, biases
exist in the fossil record: different environments are more favourable to the
preservation of different types of organism or parts of organisms. [32] Further, only the
parts of organisms that were already mineralised are usually preserved, such as the
shells of molluscs. Since most animal species are soft-bodied, they decay before
they can become fossilised. As a result, although 30-plus phyla of living animals are
known, two-thirds have never been found as fossils. [21]

This Marrella specimen illustrates how clear and detailed the fossils from the Burgess
Shale Lagerstätte are.

The Cambrian fossil record includes an unusually high number of lagerstätten, which
preserve soft tissues. These allow paleontologists to examine the internal anatomy
of animals, which in other sediments are only represented by shells, spines, claws,
etc. – if they are preserved at all. The most significant Cambrian lagerstätten are the
early Cambrian Maotianshan shale beds of Chengjiang (Yunnan, China) and Sirius
Passet (Greenland);[33] the middle Cambrian Burgess Shale (British
Columbia, Canada);[34] and the late Cambrian Orsten (Sweden) fossil beds.
While lagerstätten preserve far more than the conventional fossil record, they are far
from complete. Because lagerstätten are restricted to a narrow range of
environments (where soft-bodied organisms can be preserved very quickly, e.g. by
mudslides), most animals are probably not represented; further, the exceptional
conditions that create lagerstätten probably do not represent normal living
conditions.[35] In addition, the known Cambrian lagerstätten are rare and difficult to
date, while Precambrian lagerstätten have yet to be studied in detail.
The sparseness of the fossil record means that organisms usually exist long before
they are found in the fossil record – this is known as the Signor–Lipps effect.[36]
In 2019, a "stunning" find of lagerstätten, known as the Qingjiang biota, was reported
from the Danshui river in Hubei province, China. More than 20,000 fossil specimens
were collected, including many soft bodied animals such as jellyfish, sea anemones
and worms, as well as sponges, arthropods and algae. In some specimens the
internal body structures were sufficiently preserved that soft tissues, including
muscles, gills, mouths, guts and eyes, can be seen. The remains were dated to
around 518 Mya and around half of the species identified at the time of reporting
were previously unknown.[37][38][39]
Trace fossils[edit]

Rusophycus and other trace fossils from the Gog Group, Middle Cambrian, Lake Louise, Alberta, Canada

Trace fossils consist mainly of tracks and burrows, but also

include coprolites (fossil feces) and marks left by feeding.[40][41] Trace fossils are
particularly significant because they represent a data source that is not limited to
animals with easily fossilized hard parts, and reflects organisms' behaviour. Also,
many traces date from significantly earlier than the body fossils of animals that are
thought to have been capable of making them. [42] While exact assignment of trace
fossils to their makers is generally impossible, traces may, for example, provide the
earliest physical evidence of the appearance of moderately complex animals
(comparable to earthworms).[41]
Geochemical observations[edit]
Main article: Early Cambrian geochemical fluctuations
Several chemical markers indicate a drastic change in the environment around the
start of the Cambrian. The markers are consistent with a mass extinction, [43][44] or with
a massive warming resulting from the release of methane ice.[45] Such changes may
reflect a cause of the Cambrian explosion, although they may also have resulted
from an increased level of biological activity – a possible result of the explosion.
[45]
 Despite these uncertainties, the geochemical evidence helps by making scientists
focus on theories that are consistent with at least one of the likely environmental
changes.
Phylogenetic techniques[edit]
Cladistics is a technique for working out the "family tree" of a set of organisms. It
works by the logic that, if groups B and C have more similarities to each other than
either has to group A, then B and C are more closely related to each other than
either is to A. Characteristics that are compared may be anatomical, such as the
presence of a notochord, or molecular, by comparing sequences of DNA or protein.
The result of a successful analysis is a hierarchy of clades – groups whose members
are believed to share a common ancestor. The cladistic technique is sometimes
problematic, as some features, such as wings or camera eyes, evolved more than
once, convergently – this must be taken into account in analyses.
From the relationships, it may be possible to constrain the date that lineages first
appeared. For instance, if fossils of B or C date to X million years ago and the
calculated "family tree" says A was an ancestor of B and C, then A must have
evolved more than X million years ago.
It is also possible to estimate how long ago two living clades diverged – i.e. about
how long ago their last common ancestor must have lived  – by assuming that
DNA mutations accumulate at a constant rate. These "molecular clocks", however,
are fallible, and provide only a very approximate timing: they are not sufficiently
precise and reliable for estimating when the groups that feature in the Cambrian
explosion first evolved,[46] and estimates produced by different techniques vary by a
factor of two.[47] However, the clocks can give an indication of branching rate, and
when combined with the constraints of the fossil record, recent clocks suggest a
sustained period of diversification through the Ediacaran and Cambrian. [48]

Explanation of key scientific terms[edit]

Stem groups[49]

 —= Lines of descent
    = Basal node

    = Crown node

    = Total group

    = Crown group

    = Stem group

Phylum[edit]
A phylum is the highest level in the Linnaean system for classifying organisms. Phyla
can be thought of as groupings of animals based on general body plan. [50] Despite the
seemingly different external appearances of organisms, they are classified into phyla
based on their internal and developmental organizations. [51] For example, despite
their obvious differences, spiders and barnacles both belong to the phylum
Arthropoda, but earthworms and tapeworms, although similar in shape, belong to
different phyla. As chemical and genetic testing becomes more accurate, previously
hypothesised phyla are often entirely reworked.
A phylum is not a fundamental division of nature, such as the difference
between electrons and protons. It is simply a very high-level grouping in
a classification system created to describe all currently living organisms. This system
is imperfect, even for modern animals: different books quote different numbers of
phyla, mainly because they disagree about the classification of a huge number of
worm-like species. As it is based on living organisms, it accommodates extinct
organisms poorly, if at all.[21][52]
Stem group[edit]
The concept of stem groups was introduced to cover evolutionary "aunts" and
"cousins" of living groups, and have been hypothesized based on this scientific
theory. A crown group is a group of closely related living animals plus their last
common ancestor plus all its descendants. A stem group is a set of offshoots from
the lineage at a point earlier than the last common ancestor of the crown group; it is
a relative concept, for example tardigrades are living animals that form a crown
group in their own right, but Budd (1996) regarded them as also being a stem group
relative to the arthropods.[49][53]

Skin
(ectoderm)
Muscle
(mesoderm)
Coelom
Internal
organ
Membrane
(peritoneum)
Gut
(endoderm)
A coelomate animal is basically a set of concentric tubes, with a gap between the gut and the outer tubes.

Triploblastic[edit]
The term Triploblastic means consisting of three layers, which are formed in
the embryo, quite early in the animal's development from a single-celled egg to a
larva or juvenile form. The innermost layer forms the digestive tract (gut); the
outermost forms skin; and the middle one forms muscles and all the internal organs
except the digestive system. Most types of living animal are triploblastic – the best-
known exceptions are Porifera (sponges) and Cnidaria (jellyfish, sea anemones,
etc.).
Bilaterian[edit]
The bilaterians are animals that have right and left sides at some point in their life
histories. This implies that they have top and bottom surfaces and, importantly,
distinct front and back ends. All known bilaterian animals are triploblastic, and all
known triploblastic animals are bilaterian. Living echinoderms (sea stars, sea
urchins, sea cucumbers, etc.) 'look' radially symmetrical (like wheels) rather than
bilaterian, but their larvae exhibit bilateral symmetry and some of the earliest
echinoderms may have been bilaterally symmetrical. [54] Porifera and Cnidaria are
radially symmetrical, not bilaterian, and not triploblastic.
Coelomate[edit]
The term Coelomate means having a body cavity (coelom) containing the internal
organs. Most of the phyla featured in the debate about the Cambrian explosion [clarification
needed]
 are coelomates: arthropods, annelid worms, molluscs, echinoderms,
and chordates – the noncoelomate priapulids are an important exception. All known
coelomate animals are triploblastic bilaterians, but some triploblastic bilaterian
animals do not have a coelom – for example flatworms, whose organs are
surrounded by unspecialized tissues.

Precambrian life[edit]
Phylogenetic analysis has been used to support the view that during the Cambrian
explosion, metazoans (multi-celled animals) evolved monophyletically from a single
common ancestor: flagellated colonial protists similar to modern choanoflagellates.
[citation needed]
Evidence of animals around 1 billion years ago[edit]
Further information: Acritarch and Stromatolite

Stromatolites (Pika Formation, Middle Cambrian) near Helen Lake, Banff National Park, Canada

Modern stromatolites in Hamelin Pool Marine Nature Reserve, Western Australia

Changes in the abundance and diversity of some types of fossil have been
interpreted as evidence for "attacks" by animals or other organisms. Stromatolites,
stubby pillars built by colonies of microorganisms, are a major constituent of the
fossil record from about 2,700 million years ago, but their abundance and diversity
declined steeply after about 1,250 million years ago. This decline has been attributed
to disruption by grazing and burrowing animals. [27][28][55]
Precambrian marine diversity was dominated by small fossils known as acritarchs.
This term describes almost any small organic walled fossil – from the egg cases of
small metazoans to resting cysts of many different kinds of green algae. After
appearing around 2,000 million years ago, acritarchs underwent a boom
around 1,000 million years ago, increasing in abundance, diversity, size, complexity
of shape, and especially size and number of spines. Their increasingly spiny forms in
the last 1 billion years may indicate an increased need for defence against predation.
Other groups of small organisms from the Neoproterozoic era also show signs of
antipredator defenses.[55] A consideration of taxon longevity appears to support an
increase in predation pressure around this time. [56] In general, the fossil record shows
a very slow appearance of these lifeforms in the Precambrian, with many
cyanobacterial species making up much of the underlying sediment. [57]
Fossils of the Doushantuo formation[edit]
Main article: Doushantuo formation
The layers of the Doushantuo formation from around 580 million year old[58] harbour
microscopic fossils that may represent early bilaterians. Some have been described
as animal embryos and eggs, although some may represent the remains of giant
bacteria.[59] Another fossil, Vernanimalcula, has been interpreted as
a coelomate bilaterian,[60] but may simply be an infilled bubble.[61]
These fossils form the earliest hard-and-fast evidence of animals, as opposed to
other predators.[59][62]
Burrows[edit]
Main article: Cambrian substrate revolution

An Ediacaran trace fossil, made when an organism burrowed below a microbial mat.

The traces of organisms moving on and directly underneath the microbial mats that
covered the Ediacaran sea floor are preserved from the Ediacaran period,
about 565 million years ago.[c] They were probably made by organisms
resembling earthworms in shape, size, and how they moved. The burrow-makers
have never been found preserved, but, because they would need a head and a tail,
the burrowers probably had bilateral symmetry – which would in all probability make
them bilaterian animals.[65] They fed above the sediment surface, but were forced to
burrow to avoid predators.[66]
Around the start of the Cambrian (about 542 million years ago), many new types of
traces first appear, including well-known vertical burrows such
as Diplocraterion and Skolithos, and traces normally attributed to arthropods, such
as Cruziana and Rusophycus. The vertical burrows indicate that worm-like animals
acquired new behaviours, and possibly new physical capabilities. Some Cambrian
trace fossils indicate that their makers possessed hard exoskeletons, although they
were not necessarily mineralised.[64]
Burrows provide firm evidence of complex organisms; they are also much more
readily preserved than body fossils, to the extent that the absence of trace fossils
has been used to imply the genuine absence of large, motile, bottom-dwelling
organisms.[citation needed] They provide a further line of evidence to show that the Cambrian
explosion represents a real diversification, and is not a preservational artefact. [67]
This new habit changed the seafloor's geochemistry, and led to decreased oxygen in
the ocean and increased CO2-levels in the seas and the atmosphere, resulting in
global warming for tens of millions years, and could be responsible for mass
extinctions.[68] But as burrowing became established, it allowed an explosion of its
own, for as burrowers disturbed the sea floor, they aerated it, mixing oxygen into the
toxic muds. This made the bottom sediments more hospitable, and allowed a wider
range of organisms to inhabit them – creating new niches and the scope for higher
diversity.[67]
Ediacaran organisms[edit]
Dickinsonia costata, an Ediacaran organism of unknown affinity, with a quilted appearance

Main articles: Ediacara biota, Cloudinid, Kimberella, and Spriggina

At the start of the Ediacaran period, much of the acritarch fauna, which had
remained relatively unchanged for hundreds of millions of years, became extinct, to
be replaced with a range of new, larger species, which would prove far more
ephemeral.[57] This radiation, the first in the fossil record,[57] is followed soon after by an
array of unfamiliar, large, fossils dubbed the Ediacara biota, [69] which flourished for 40
million years until the start of the Cambrian.[70] Most of this "Ediacara biota" were at
least a few centimeters long, significantly larger than any earlier fossils. The
organisms form three distinct assemblages, increasing in size and complexity as
time progressed.[71]
Many of these organisms were quite unlike anything that appeared before or since,
resembling discs, mud-filled bags, or quilted mattresses – one palæontologist
proposed that the strangest organisms should be classified as a separate kingdom,
Vendozoa.[72]

Fossil of Kimberella, a triploblastic bilaterian, and possibly a mollusc

At least some may have been early forms of the phyla at the heart of the "Cambrian
explosion" debate,[clarification needed] having been interpreted as early molluscs (Kimberella),[29]
[73]
 echinoderms (Arkarua);[74] and arthropods (Spriggina,[75] Parvancorina,[76] Yilingia).
Still, debate exists about the classification of these specimens, mainly because the
diagnostic features that allow taxonomists to classify more recent organisms, such
as similarities to living organisms, are generally absent in the ediacarans. [77] However,
there seems little doubt that Kimberella was at least a triploblastic bilaterian animal.
[77]
 These organisms are central to the debate about how abrupt the Cambrian
explosion was.[citation needed] If some were early members of the animal phyla seen today,
the "explosion" looks a lot less sudden than if all these organisms represent an
unrelated "experiment", and were replaced by the animal kingdom fairly soon
thereafter (40M years is "soon" by evolutionary and geological standards).
Beck Spring Dolomite[edit]
Paul Knauth, a geologist at Arizona State University, maintains that
photosynthesizing organisms such as algae may have grown over a 750- to 800-
million-year-old formation in Death Valley known as the Beck Spring Dolomite. In the
early 1990s, samples from this 1,000-foot thick layer of dolomite revealed that the
region housed flourishing mats of photosynthesizing, unicellular life forms which
antedated the Cambrian explosion.
Microfossils have been unearthed from holes riddling the otherwise barren surface of
the dolomite. These geochemical and microfossil findings support the idea that
during the Precambrian period, complex life evolved both in the oceans and on land.
Knauth contends that animals may well have had their origins in freshwater lakes
and streams, and not in the oceans.
Some 30 years later, a number of studies have documented an abundance of
geochemical and microfossil evidence showing that life covered the continents as far
back as 2.2 billion years ago. Many paleobiologists now accept the idea that simple
life forms existed on land during the Precambrian, but are opposed to the more
radical idea that multicellular life thrived on land more than 600 million years ago. [78]

Ediacaran–Early Cambrian skeletonisation[edit]

The first Ediacaran and lowest Cambrian (Nemakit-Daldynian) skeletal fossils
represent tubes and problematic sponge spicules. [79] The oldest sponge spicules are
monaxon siliceous, aged around 580 million years ago, known from the Doushantou
Formation in China and from deposits of the same age in Mongolia, although the
interpretation of these fossils as spicules has been challenged. [80] In the late
Ediacaran-lowest Cambrian, numerous tube dwellings of enigmatic organisms
appeared. It was organic-walled tubes (e.g. Saarina) and chitinous tubes of the
sabelliditids (e.g. Sokoloviina, Sabellidites, Paleolina)[81][82] that prospered up to the
beginning of the Tommotian. The mineralized tubes
of Cloudina, Namacalathus, Sinotubulites, and a dozen more of the other organisms
from carbonate rocks formed near the end of the Ediacaran period
from 549 to 542 million years ago, as well as the triradially symmetrical mineralized
tubes of anabaritids (e.g. Anabarites, Cambrotubulus) from uppermost Ediacaran
and lower Cambrian.[83] Ediacaran mineralized tubes are often found in carbonates of
the stromatolite reefs and thrombolites,[84][85] i.e. they could live in an environment
adverse to the majority of animals.
Although they are as hard to classify as most other Ediacaran organisms, they are
important in two other ways. First, they are the earliest known calcifying organisms
(organisms that built shells from calcium carbonate).[85][86][87] Secondly, these tubes are
a device to rise over a substrate and competitors for effective feeding and, to a
lesser degree, they serve as armor for protection against predators and adverse
conditions of environment. Some Cloudina fossils show small holes in shells. The
holes possibly are evidence of boring by predators sufficiently advanced to penetrate
shells.[88] A possible "evolutionary arms race" between predators and prey is one of
the hypotheses that attempt to explain the Cambrian explosion. [55]
In the lowest Cambrian, the stromatolites were decimated. This allowed animals to
begin colonization of warm-water pools with carbonate sedimentation. At first, it
was anabaritids and Protohertzina (the fossilized grasping spines of chaetognaths)
fossils. Such mineral skeletons as shells, sclerites, thorns, and plates appeared in
uppermost Nemakit-Daldynian; they were the earliest species
of halkierids, gastropods, hyoliths and other rare organisms. The beginning of
the Tommotian has historically been understood to mark an explosive increase of the
number and variety of fossils of molluscs, hyoliths, and sponges, along with a rich
complex of skeletal elements of unknown animals, the
first archaeocyathids, brachiopods, tommotiids, and others.[89][90][91][92] Also soft-
bodied extant phyla such as comb jellies, scalidophorans, entoproctans, horseshoe
worms and lobopodians had armored forms.[93] This sudden increase is partially an
artefact of missing strata at the Tommotian type section, and most of this fauna in
fact began to diversify in a series of pulses through the Nemakit-Daldynian and into
the Tommotian.[94]
Some animals may already have had sclerites, thorns, and plates in the Ediacaran
(e.g. Kimberella had hard sclerites, probably of carbonate), but thin carbonate
skeletons cannot be fossilized in siliciclastic deposits.[95] Older (~750 Ma) fossils
indicate that mineralization long preceded the Cambrian, probably defending small
photosynthetic algae from single-celled eukaryotic predators. [96][97]

Cambrian life[edit]
Trace fossils[edit]
Trace fossils (burrows, etc.) are a reliable indicator of what life was around, and
indicate a diversification of life around the start of the Cambrian, with the freshwater
realm colonized by animals almost as quickly as the oceans. [98]
Small shelly fauna[edit]
Main article: Small shelly fauna
Fossils known as "small shelly fauna" have been found in many parts on the world,
and date from just before the Cambrian to about 10 million years after the start of the
Cambrian (the Nemakit-Daldynian and Tommotian ages; see timeline). These are a
very mixed collection of fossils: spines, sclerites (armor plates),
tubes, archeocyathids (sponge-like animals), and small shells very like those
of brachiopods and snail-like molluscs – but all tiny, mostly 1 to 2 mm long.[99]
While small, these fossils are far more common than complete fossils of the
organisms that produced them; crucially, they cover the window from the start of the
Cambrian to the first lagerstätten: a period of time otherwise lacking in fossils.
Hence, they supplement the conventional fossil record and allow the fossil ranges of
many groups to be extended.
Early Cambrian trilobites and echinoderms[edit]
A fossilized trilobite, an ancient type of arthropod: This specimen, from the Burgess Shale, preserves "soft
parts" – the antennae and legs.

The earliest trilobite fossils are about 530 million years old, but the class was already
quite diverse and worldwide, suggesting they had been around for quite some time.
[100]
 The fossil record of trilobites began with the appearance of trilobites with mineral
exoskeletons – not from the time of their origin.
The earliest generally accepted echinoderm fossils appeared a little bit later, in the
Late Atdabanian; unlike modern echinoderms, these early Cambrian echinoderms
were not all radially symmetrical.[101]
These provide firm data points for the "end" of the explosion, or at least indications
that the crown groups of modern phyla were represented.
Burgess Shale type faunas[edit]
Main article: Burgess Shale type preservation
The Burgess Shale and similar lagerstätten preserve the soft parts of organisms,
which provide a wealth of data to aid in the classification of enigmatic fossils. It often
preserved complete specimens of organisms only otherwise known from dispersed
parts, such as loose scales or isolated mouthparts. Further, the majority of
organisms and taxa in these horizons are entirely soft-bodied, hence absent from the
rest of the fossil record.[102] Since a large part of the ecosystem is preserved, the
ecology of the community can also be tentatively reconstructed. [verification needed] However,
the assemblages may represent a "museum": a deep-water ecosystem that is
evolutionarily "behind" the rapidly diversifying fauna of shallower waters. [103]
Because the lagerstätten provide a mode and quality of preservation that is virtually
absent outside of the Cambrian, many organisms appear completely different from
anything known from the conventional fossil record. This led early workers in the field
to attempt to shoehorn the organisms into extant phyla; the shortcomings of this
approach led later workers to erect a multitude of new phyla to accommodate all the
oddballs. It has since been realised that most oddballs diverged from lineages before
they established the phyla known today[clarification needed] – slightly different designs, which
were fated to perish rather than flourish into phyla, as their cousin lineages did.
The preservational mode is rare in the preceding Ediacaran period, but those
assemblages known show no trace of animal life – perhaps implying a genuine
absence of macroscopic metazoans.[104]
Early Cambrian crustaceans[edit]
Further information: Orsten
Crustaceans, one of the four great modern groups of arthropods, are very rare
throughout the Cambrian. Convincing crustaceans were once thought to be common
in Burgess Shale-type biotas, but none of these individuals can be shown to fall into
the crown group of "true crustaceans".[105] The Cambrian record of crown-group
crustaceans comes from microfossils. The Swedish Orsten horizons contain later
Cambrian crustaceans, but only organisms smaller than 2 mm are preserved. This
restricts the data set to juveniles and miniaturised adults.
A more informative data source is the organic microfossils of the Mount Cap
formation, Mackenzie Mountains, Canada. This late Early Cambrian assemblage
(510 to 515 million years ago) consists of microscopic fragments of arthropods'
cuticle, which is left behind when the rock is dissolved with hydrofluoric acid. The
diversity of this assemblage is similar to that of modern crustacean faunas. Analysis
of fragments of feeding machinery found in the formation shows that it was adapted
to feed in a very precise and refined fashion. This contrasts with most other early
Cambrian arthropods, which fed messily by shovelling anything they could get their
feeding appendages on into their mouths. This sophisticated and specialised feeding
machinery belonged to a large (about 30 cm)[106] organism, and would have provided
great potential for diversification; specialised feeding apparatus allows a number of
different approaches to feeding and development, and creates a number of different
approaches to avoid being eaten.[105]
Early Ordovician radiation[edit]
Main article: Ordovician radiation
After an extinction at the Cambrian–Ordovician boundary, another radiation
occurred, which established the taxa that would dominate the Palaeozoic. [107]
During this radiation, the total number of orders doubled, and families tripled,
[107]
 increasing marine diversity to levels typical of the Palaeozoic, [45] and disparity to
levels approximately equivalent to today's. [11]

Stages[edit]
The event lasted for about the next 20[5][108]–25[109][110] million years, and its elevated rates
of evolution had ended by the base of Cambrian Series 2, 521 million years ago,
coincident with the first trilobites in the fossil record. [111] Different authors break the
explosion down into stages in different ways.
Ed Landing recognizes three stages: Stage 1, spanning the Ediacaran-Cambrian
boundary, corresponds to a diversification of biomineralizing animals and of deep
and complex burrows; Stage 2, corresponding to the radiation of molluscs and stem-
group Brachiopods (hyoliths and tommotiids), which apparently arose in intertidal
waters; and Stage 3, seeing the Atdabanian diversification of trilobites in deeper
waters, but little change in the intertidal realm. [112]
Graham Budd synthesises various schemes to produce a compatible view of the
SSF record of the Cambrian explosion, divided slightly differently into four intervals:
a "Tube world", lasting from 550 to 536 million years ago, spanning the Ediacaran-
Cambrian boundary, dominated by Cloudina, Namacalathus and pseudoconodont-
type elements; a "Sclerite world", seeing the rise of halkieriids, tommotiids, and
hyoliths, lasting to the end of the Fortunian (c. 525 Ma); a brachiopod world, perhaps
corresponding to the as yet unratified Cambrian Stage 2; and Trilobite World, kicking
off in Stage 3.[113]
Complementary to the shelly fossil record, trace fossils can be divided into five
subdivisions: "Flat world" (late Ediacaran), with traces restricted to the sediment
surface; Protreozoic III (after Jensen), with increasing complexity; pedum world,
initiated at the base of the Cambrian with the base of the T.pedum zone (see
discussion at Cambrian#Dating the Cambrian); Rusophycus world,
spanning 536 to 521 million years ago and thus corresponding exactly to the periods
of Sclerite World and Brachiopod World under the SSF paradigm;
and Cruziana world, with an obvious correspondence to Trilobite World. [113]

Validity[edit]
There is strong evidence for species of Cnidaria and Porifera existing in
the Ediacaran[114] and possible members of Porifera even before that during
the Cryogenian.[115] Bryozoans don't appear in the fossil record until after the
Cambrian, in the Lower Ordovician.[116]
The fossil record as Darwin knew it seemed to suggest that the major metazoan
groups appeared in a few million years of the early to mid-Cambrian, and even in the
1980s, this still appeared to be the case. [24][25]
However, evidence of Precambrian Metazoa is gradually accumulating. If the
Ediacaran Kimberella was a mollusc-like protostome (one of the two main groups
of coelomates),[29][73] the protostome and deuterostome lineages must have split
significantly before 550 million years ago (deuterostomes are the other main group of
coelomates).[117] Even if it is not a protostome, it is widely accepted as a bilaterian. [77]
[117]
 Since fossils of rather modern-looking cnidarians (jellyfish-like organisms) have
been found in the Doushantuo lagerstätte, the cnidarian and bilaterian lineages must
have diverged well over 580 million years ago.[117]
Trace fossils[71] and predatory borings in Cloudina shells provide further evidence of
Ediacaran animals.[118] Some fossils from the Doushantuo formation have been
interpreted as embryos and one (Vernanimalcula) as a bilaterian coelomate,
although these interpretations are not universally accepted. [60][61][119] Earlier still,
predatory pressure has acted on stromatolites and acritarchs since
around 1,250 million years ago.[55]
Some say that the evolutionary change was accelerated by an order of magnitude,
[d]
 but the presence of Precambrian animals somewhat dampens the "bang" of the
explosion; not only was the appearance of animals gradual, but their evolutionary
radiation ("diversification") may also not have been as rapid as once thought. Indeed,
statistical analysis shows that the Cambrian explosion was no faster than any of the
other radiations in animals' history.[e] However, it does seem that some innovations
linked to the explosion – such as resistant armour – only evolved once in the animal
lineage; this makes a lengthy Precambrian animal lineage harder to defend.
[121]
 Further, the conventional view that all the phyla arose in the Cambrian is flawed;
while the phyla may have diversified in this time period, representatives of the crown
groups of many phyla do not appear until much later in the Phanerozoic. [12] Further,
the mineralised phyla that form the basis of the fossil record may not be
representative of other phyla, since most mineralised phyla originated in
a benthic setting. The fossil record is consistent with a Cambrian explosion that was
limited to the benthos, with pelagic phyla evolving much later. [12]
Ecological complexity among marine animals increased in the Cambrian, as well
later in the Ordovician.[11] However, recent research has overthrown the once-popular
idea that disparity was exceptionally high throughout the Cambrian, before
subsequently decreasing.[122] In fact, disparity remains relatively low throughout the
Cambrian, with modern levels of disparity only attained after the early Ordovician
radiation.[11]
The diversity of many Cambrian assemblages is similar to today's, [123][105] and at a high
(class/phylum) level, diversity is thought by some to have risen relatively smoothly
through the Cambrian, stabilizing somewhat in the Ordovician. [124] This interpretation,
however, glosses over the astonishing and fundamental pattern of
basal polytomy and phylogenetic telescoping at or near the Cambrian boundary, as
seen in most major animal lineages.[125] Thus Harry Blackmore Whittington's
questions regarding the abrupt nature of the Cambrian explosion remain, and have
yet to be satisfactorily answered.[126]
The Cambrian explosion as survivorship bias[edit]
Budd and Mann[127] suggested that the Cambrian explosion was the result of a type
of survivorship bias called the "Push of the past". As groups at their origin tend to go
extinct, it follows that any long-lived group would have experienced an unusually
rapid rate of diversification early on, creating the illusion of a general speed-up in
diversification rates. However, rates of diversification could remain at background
levels and still generate this sort of effect in the surviving lineages.

Possible causes[edit]
Despite the evidence that moderately complex animals (triploblastic bilaterians)
existed before and possibly long before the start of the Cambrian, it seems that the
pace of evolution was exceptionally fast in the early Cambrian. Possible explanations
for this fall into three broad categories: environmental, developmental, and ecological
changes. Any explanation must explain both the timing and magnitude of the
explosion.
Changes in the environment[edit]
Increase in oxygen levels[edit]
Earth's earliest atmosphere contained no free oxygen (O2); the oxygen that animals
breathe today, both in the air and dissolved in water, is the product of billions of
years of photosynthesis. Cyanobacteria were the first organisms to evolve the ability
to photosynthesize, introducing a steady supply of oxygen into the environment.
[128]
 Initially, oxygen levels did not increase substantially in the atmosphere. [129] The
oxygen quickly reacted with iron and other minerals in the surrounding rock and
ocean water. Once a saturation point was reached for the reactions in rock and
water, oxygen was able to exist as a gas in its diatomic form. Oxygen levels in the
atmosphere increased substantially afterward. [130] As a general trend,
the concentration of oxygen in the atmosphere has risen gradually over about the
last 2.5 billion years.[21]
Oxygen levels seem to have a positive correlation with diversity in eukaryotes well
before the Cambrian period.[131] The last common ancestor of all extant eukaryotes is
thought to have lived around 1.8 billion years ago. Around 800 million years ago,
there was a notable increase in the complexity and number of eukaryotes species in
the fossil record.[131] Before the spike in diversity, eukaryotes are thought to have lived
in highly sulfuric environments. Sulfide interferes with mitochondrial function in
aerobic organisms, limiting the amount of oxygen that could be used to drive
metabolism. Oceanic sulfide levels decreased around 800 million years ago, which
supports the importance of oxygen in eukaryotic diversity. [131]
The shortage of oxygen might well have prevented the rise of large, complex
animals. The amount of oxygen an animal can absorb is largely determined by the
area of its oxygen-absorbing surfaces (lungs and gills in the most complex animals;
the skin in less complex ones); but, the amount needed is determined by its volume,
which grows faster than the oxygen-absorbing area if an animal's size increases
equally in all directions. An increase in the concentration of oxygen in air or water
would increase the size to which an organism could grow without its tissues
becoming starved of oxygen. However, members of the Ediacara biota reached
metres in length tens of millions of years before the Cambrian explosion. [43] Other
metabolic functions may have been inhibited by lack of oxygen, for example the
construction of tissue such as collagen, required for the construction of complex
structures,[132] or to form molecules for the construction of a hard exoskeleton.
[133]
 However, animals were not affected when similar oceanographic conditions
occurred in the Phanerozoic; there is no convincing correlation between oxygen
levels and evolution, so oxygen may have been no more a prerequisite to complex
life than liquid water or primary productivity.[134]
Ozone formation[edit]
The amount of ozone (O3) required to shield Earth from biologically lethal UV
radiation, wavelengths from 200 to 300 nanometers (nm), is believed to have been in
existence around the Cambrian explosion.[135] The presence of the ozone layer may
have enabled the development of complex life and life on land, as opposed to life
being restricted to the water.
Snowball Earth[edit]
Main article: Snowball Earth
In the late Neoproterozoic (extending into the early Ediacaran period), the Earth
suffered massive glaciations in which most of its surface was covered by ice. This
may have caused a mass extinction, creating a genetic bottleneck; the resulting
diversification may have given rise to the Ediacara biota, which appears soon after
the last "Snowball Earth" episode.[136] However, the snowball episodes occurred a
long time before the start of the Cambrian, and it is difficult to see how so much
diversity could have been caused by even a series of bottlenecks; [45] the cold periods
may even have delayed the evolution of large size organisms.[55]
Increase in the calcium concentration of the Cambrian seawater [edit]
Newer research suggests that volcanically active midocean ridges caused a massive
and sudden surge of the calcium concentration in the oceans, making it possible for
marine organisms to build skeletons and hard body parts. [137] Alternatively a high
influx of ions could have been provided by the widespread erosion that produced
Powell's Great Unconformity.[138]
An increase of calcium may also have been caused by erosion of
the Transgondwanan Supermountain that existed at the time of the explosion. The
roots of the mountain are preserved in present-day East Africa as an orogen.[139]
Developmental explanations[edit]
Further information: Evolutionary developmental biology
A range of theories are based on the concept that minor modifications to animals'
development as they grow from embryo to adult may have been able to cause very
large changes in the final adult form. The Hox genes, for example, control which
organs individual regions of an embryo will develop into. For instance, if a
certain Hox gene is expressed, a region will develop into a limb; if a different Hox
gene is expressed in that region (a minor change), it could develop into an eye
instead (a phenotypically major change).
Such a system allows a large range of disparity to appear from a limited set of
genes, but such theories linking this with the explosion struggle to explain why the
origin of such a development system should by itself lead to increased diversity or
disparity. Evidence of Precambrian metazoans[45] combines with molecular data[140] to
show that much of the genetic architecture that could feasibly have played a role in
the explosion was already well established by the Cambrian.
This apparent paradox is addressed in a theory that focuses on the physics of
development. It is proposed that the emergence of simple multicellular forms
provided a changed context and spatial scale in which novel physical processes and
effects were mobilized by the products of genes that had previously evolved to serve
unicellular functions. Morphological complexity (layers, segments, lumens,
appendages) arose, in this view, by self-organization.[141]
Horizontal gene transfer has also been identified as a possible factor in the rapid
acquisition of the biochemical capability of biomineralization among organisms
during this period, based on evidence that the gene for a critical protein in the
process was originally transferred from a bacterium into sponges. [142]
Ecological explanations[edit]
These focus on the interactions between different types of organism. Some of these
hypotheses deal with changes in the food chain; some suggest arms races between
predators and prey, and others focus on the more general mechanisms
of coevolution. Such theories are well suited to explaining why there was a rapid
increase in both disparity and diversity, but they do not explain why the "explosion"
happened when it did.[45]
End-Ediacaran mass extinction[edit]
Main article: End-Ediacaran extinction
Evidence for such an extinction includes the disappearance from the fossil record of
the Ediacara biota and shelly fossils such as Cloudina, and the accompanying
perturbation in the δ13C record. It is suspected that several global anoxic events were
responsible for the extinction.[143][144]
Mass extinctions are often followed by adaptive radiations as existing clades expand
to occupy the ecospace emptied by the extinction. However, once the dust had
settled, overall disparity and diversity returned to the pre-extinction level in each of
the Phanerozoic extinctions.[45]
Anoxia[edit]
The late Ediacaran oceans appears to have suffered from an anoxia that covered
much of the seafloor, which would have given mobile animals able to seek out more
oxygen-rich environments an advantage over sessile forms of life. [145]
Evolution of eyes[edit]
Main article: Evolution of the eye
Andrew Parker has proposed that predator-prey relationships changed dramatically
after eyesight evolved. Prior to that time, hunting and evading were both close-range
affairs – smell, vibration, and touch were the only senses used. When predators
could see their prey from a distance, new defensive strategies were needed. Armor,
spines, and similar defenses may also have evolved in response to vision. He further
observed that, where animals lose vision in unlighted environments such as caves,
diversity of animal forms tends to decrease.[146] Nevertheless, many scientists doubt
that vision could have caused the explosion. Eyes may well have evolved long
before the start of the Cambrian.[147] It is also difficult to understand why the evolution
of eyesight would have caused an explosion, since other senses, such as smell and
pressure detection, can detect things at a greater distance in the sea than sight can;
but the appearance of these other senses apparently did not cause an evolutionary
explosion.[45]
Arms races between predators and prey[edit]
The ability to avoid or recover from predation often makes the difference between life
and death, and is therefore one of the strongest components of natural selection.
The pressure to adapt is stronger on the prey than on the predator: if the predator
fails to win a contest, it loses a meal; if the prey is the loser, it loses its life. [148]
But, there is evidence that predation was rife long before the start of the Cambrian,
for example in the increasingly spiny forms of acritarchs, the holes drilled
in Cloudina shells, and traces of burrowing to avoid predators. Hence, it is unlikely
that the appearance of predation was the trigger for the Cambrian "explosion",
although it may well have exhibited a strong influence on the body forms that the
"explosion" produced.[55] However, the intensity of predation does appear to have
increased dramatically during the Cambrian [149] as new predatory "tactics" (such as
shell-crushing) emerged.[150] This rise of predation during the Cambrian was confirmed
by the temporal pattern of the median predator ratio at the scale of genus, in fossil
communities covering the Cambrian and Ordovician periods, but this pattern is not
correlated to diversification rate.[151] This lack of correlation between predator ratio and
diversification over the Cambrian and Ordovician suggests that predators did not
trigger the large evolutionary radiation of animals during this interval. Thus the role of
predators as triggerers of diversification may have been limited to the very beginning
of the "Cambrian explosion".[151]
Increase in size and diversity of planktonic animals [edit]
Geochemical evidence strongly indicates that the total mass of plankton has been
similar to modern levels since early in the Proterozoic. Before the start of the
Cambrian, their corpses and droppings were too small to fall quickly towards the
seabed, since their drag was about the same as their weight. This meant they were
destroyed by scavengers or by chemical processes before they reached the sea
floor.[35]
Mesozooplankton are plankton of a larger size. Early Cambrian
specimens filtered microscopic plankton from the seawater. These larger organisms
would have produced droppings and ultimately corpses large enough to fall fairly
quickly. This provided a new supply of energy and nutrients to the mid-levels and
bottoms of the seas, which opened up a new range of possible ways of life. If any of
these remains sank uneaten to the sea floor they could be buried; this would have
taken some carbon out of circulation, resulting in an increase in the concentration of
breathable oxygen in the seas (carbon readily combines with oxygen).[35]
The initial herbivorous mesozooplankton were probably larvae of benthic (seafloor)
animals. A larval stage was probably an evolutionary innovation driven by the
increasing level of predation at the seafloor during the Ediacaran period.[10][152]
Metazoans have an amazing ability to increase diversity through coevolution.[57] This
means that an organism's traits can lead to traits evolving in other organisms; a
number of responses are possible, and a different species can potentially emerge
from each one. As a simple example, the evolution of predation may have caused
one organism to develop a defence, while another developed motion to flee. This
would cause the predator lineage to diverge into two species: one that was good at
chasing prey, and another that was good at breaking through defences. Actual
coevolution is somewhat more subtle, but, in this fashion, great diversity can arise:
three quarters of living species are animals, and most of the rest have formed by
coevolution with animals.[57]
Ecosystem engineering[edit]
Evolving organisms inevitably change the environment they evolve in.
The Devonian colonization of land had planet-wide consequences for sediment
cycling and ocean nutrients, and was likely linked to the Devonian mass extinction. A
similar process may have occurred on smaller scales in the oceans, with, for
example, the sponges filtering particles from the water and depositing them in the
mud in a more digestible form; or burrowing organisms making previously
unavailable resources available for other organisms. [153]
Complexity threshold[edit]
The explosion may not have been a significant evolutionary event. It may represent a
threshold being crossed: for example a threshold in genetic complexity that allowed
a vast range of morphological forms to be employed. [154] This genetic threshold may
have a correlation to the amount of oxygen available to organisms. Using oxygen for
metabolism produces much more energy than anaerobic processes. Organisms that
use more oxygen have the opportunity to produce more complex proteins, providing
a template for further evolution.[129] These proteins translate into larger, more complex
structures that allow organisms better to adapt to their environments. [155] With the help
of oxygen, genes that code for these proteins could contribute to the expression
of complex traits more efficiently. Access to a wider range of structures and functions
would allow organisms to evolve in different directions, increasing the number of
niches that could be inhabited. Furthermore, organisms had the opportunity to
become more specialized in their own niches.[155]

Uniqueness of the explosion[edit]

The "Cambrian explosion" can be viewed as two waves of metazoan expansion into
empty niches: first, a coevolutionary rise in diversity as animals explored niches on
the Ediacaran sea floor, followed by a second expansion in the early Cambrian as
they became established in the water column.[57] The rate of diversification seen in the
Cambrian phase of the explosion is unparalleled among marine animals: it affected
all metazoan clades of which Cambrian fossils have been found. Later radiations,
such as those of fish in the Silurian and Devonian periods, involved fewer taxa,
mainly with very similar body plans.[21] Although the recovery from the Permian-
Triassic extinction started with about as few animal species as the Cambrian
explosion, the recovery produced far fewer significantly new types of animals. [156]
Whatever triggered the early Cambrian diversification opened up an exceptionally
wide range of previously unavailable ecological niches. When these were all
occupied, limited space existed for such wide-ranging diversifications to occur again,
because strong competition existed in all niches and incumbents usually had the
advantage. If a wide range of empty niches had continued, clades would be able to
continue diversifying and become disparate enough for us to recognise them as
different phyla; when niches are filled, lineages will continue to resemble one another
long after they diverge, as limited opportunity exists for them to change their life-
styles and forms.[157]
There were two similar explosions in the evolution of land plants: after a cryptic
history beginning about 450 million years ago, land plants underwent a uniquely
rapid adaptive radiation during the Devonian period, about 400 million years ago.
[21]
 Furthermore, Angiosperms (flowering plants) originated and rapidly diversified
during the Cretaceous period.

See also[edit]
 Massive Australian Precambrian/Cambrian Impact
Structure

Footnotes[edit]
1. ^ This included at least animals, phytoplankton and calcimicrobes.
[10]

2. ^ At 610 million years ago, Aspidella disks appeared, but it is not

clear that these represented complex life forms.
3. ^ Older marks found in billion-year-old rocks[63] have since been
recognised as nonbiogenic.[12][64]
4. ^ As defined in terms of the extinction and origination rate of
species.[57]
5. ^ The analysis considered the bioprovinciality of trilobite lineages,
as well as their evolutionary rate.[120]

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Cambrian explosion".  Int. J. Dev. Biol.  47  (7): 685–693.  arXiv:q-
bio/0311013. Bibcode:2003q.bio....11013S.  PMID  14756344.
155. ^ Jump up to:a b Sperling, E.A.; Frieder, C.A.; Raman, A.V.; Girguis,
P.R.; Levin, L.A.; Knoll, A.H. (2013). "Oxygen, ecology, and the
Cambrian radiation of animals". PNAS.  110  (33): 13446–
13451. Bibcode:2013PNAS..11013446S. doi:10.1073/pnas.13127
78110. PMC  3746845.  PMID  23898193.
156. ^ Erwin, D.H.; Valentine, J.W.; Sepkoski, J.J. (November 1987).
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Paleozoic Versus the Mesozoic". Evolution.  41  (6): 1177–
1186.  doi:10.2307/2409086.  JSTOR  2409086.  PMID  11542112.
157. ^ Valentine, J.W. (April 1995). "Why No New Phyla after the
Cambrian? Genome and Ecospace Hypotheses
Revisited".  PALAIOS  (abstract). 10 (2): 190–
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3515182.
Further reading[edit]
 Budd, G. E.; Jensen, J. (2000). "A critical reappraisal of
the fossil record of the bilaterian phyla". Biological
Reviews. 75 (2): 253–295. doi:10.1111/j.1469-
185X.1999.tb00046.x. PMID 10881389.
 Collins, Allen G. "Metazoa: Fossil record". Retrieved
Dec. 14, 2005.
 Conway Morris, S. (1997). The Crucible of Creation: the
Burgess Shale and the rise of animals. Oxford
University Press. ISBN 0-19-286202-2.
 Conway Morris, S. (June 2006). "Darwin's dilemma: the
realities of the Cambrian 'explosion'". Philosophical
Transactions of the Royal Society B: Biological
Sciences. 361 (1470): 1069–
1083. doi:10.1098/rstb.2006.1846. ISSN 0962-8436. P
MC 1578734. PMID 16754615. An enjoyable account.
 Gould, S.J. (1989). Wonderful Life: The Burgess Shale
and the Nature of History. W.W. Norton &
Company. Bibcode:1989wlbs.book.....G.
 Kennedy, M.; M. Droser; L. Mayer.; D. Pevear & D.
Mrofka (2006). "Clay and Atmospheric
Oxygen". Science. 311 (5766):
1341. doi:10.1126/science.311.5766.1341c.
 Knoll, A.H.; Carroll, S.B. (1999-06-25). "Early Animal
Evolution: Emerging Views from Comparative Biology
and Geology". Science. 284 (5423): 2129–
37. doi:10.1126/science.284.5423.2129. PMID 103818
72. S2CID 8908451.
 Markov, Alexander V.; Korotayev, Andrey V. (2007).
"Phanerozoic marine biodiversity follows a hyperbolic
trend". Palaeoworld. 16 (4): 311–
318. doi:10.1016/j.palwor.2007.01.002.
 Montenari, M.; Leppig, U. (2003). "The Acritarcha: their
classification morphology, ultrastructure and
palaeoecological/palaeogeographical
distribution". Paläontologische Zeitschrift. 77: 173–
194. doi:10.1007/bf03004567.
 Wang, D. Y.-C.; S. Kumar; S. B. Hedges (January
1999). "Divergence time estimates for the early history
of animal phyla and the origin of plants, animals and
fungi". Proceedings of the Royal Society B. 266 (1415):
163–71. doi:10.1098/rspb.1999.0617. ISSN 0962-8452. 
PMC 1689654. PMID 10097391.
 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.
 Xiao, S.; Y. Zhang & A. Knoll (January 1998). "Three-
dimensional preservation of algae and animal embryos
in a Neoproterozoic phosphorite". Nature. 391 (1): 553–
58. Bibcode:1998Natur.391..553X. doi:10.1038/35318. 
ISSN 0090-9556.
Timeline References:

 Martin, M.W; Grazhdankin, D.V; Bowring, S.A; Evans,

D.A.D; Fedonkin, M.A; Kirschvink, J.L (2000). "Age of
Neoproterozoic Bilaterian Body and Trace Fossils,
White Sea, Russia: Implications for Metazoan
Evolution". Science. 288 (5467): 841–
845. Bibcode:2000Sci...288..841M. doi:10.1126/scienc
e.288.5467.841. PMID 10797002.

External links[edit]
 The Cambrian "explosion" of metazoans and molecular
biology: would Darwin be satisfied?
 On embryos and ancestors by Stephen Jay Gould
 Conway Morris, S. (April 2000). "The Cambrian
"explosion": Slow-fuse or megatonnage?". Proceedings
of the National Academy of Sciences. 97 (9): 4426–
4429. Bibcode:2000PNAS...97.4426C. doi:10.1073/pna
s.97.9.4426. PMC 34314. PMID 10781036.
 The Cambrian Explosion – In Our Time, BBC Radio
4 broadcast, 17 February 2005
 "Burgess Shale". Virtual Museum of Canada. 2011.,
exhaustive details about the Burgess Shale, its fossils,
and its significance for the Cambrian explosion
 Utah's Cambrian life – new (2008) website with good
images of a range of Burgess-shale-type and other
Cambrian fossils
 Smithsonian National Museum
Categories: 
 Cambrian animals
 Cambrian events
 Cambrian first appearances
 Cambrian life
 Evolution
 Unsolved problems in biology
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