Doushantuo Formation
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The Doushantuo Formation (formerly transcribed as Toushantuo or Toushantou, from ) is a
geological formation A geological formation, or simply formation, is a body of rock having a consistent set of physical characteristics ( lithology) that distinguishes it from adjacent bodies of rock, and which occupies a particular position in the layers of rock exp ...
in western
Hubei Hubei (; ; alternately Hupeh) is a landlocked province of the People's Republic of China, and is part of the Central China region. The name of the province means "north of the lake", referring to its position north of Dongting Lake. The prov ...
, eastern
Guizhou Guizhou (; formerly Kweichow) is a landlocked province in the southwest region of the People's Republic of China. Its capital and largest city is Guiyang, in the center of the province. Guizhou borders the autonomous region of Guangxi to t ...
, southern
Shaanxi Shaanxi (alternatively Shensi, see #Name, § Name) is a landlocked Provinces of China, province of China. Officially part of Northwest China, it borders the province-level divisions of Shanxi (NE, E), Henan (E), Hubei (SE), Chongqing (S), Sichu ...
, central
Jiangxi Jiangxi (; ; formerly romanized as Kiangsi or Chianghsi) is a landlocked province in the east of the People's Republic of China. Its major cities include Nanchang and Jiujiang. Spanning from the banks of the Yangtze river in the north int ...
, and other localities in
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
. It is known for the fossil Lagerstätten in Zigui in Hubei,
Xiuning Xiuning County () is a county in the south of Anhui Province, People's Republic of China, under the jurisdiction of the prefecture-level city of Huangshan City. The southernmost county-level division in the province, it has a population of and an ...
in Anhui, and Weng'an in Guizhou, as one of the oldest beds to contain minutely preserved
microfossil A microfossil is a fossil that is generally between 0.001 mm and 1 mm in size, the visual study of which requires the use of light or electron microscopy. A fossil which can be studied with the naked eye or low-powered magnification, ...
s, phosphatic fossils that are so characteristic they have given their name to "
Doushantuo type preservation The preservational mode of the Doushantuo formation involves very early phosphatisation on a cellular level - with cells being replaced by phosphate before they degrade. Occurrence The mode of preservation is typically found in shallow, high energy ...
". The formation, whose deposits date back to the Early and Middle Ediacaran, is of particular interest because it covers the poorly understood interval of time between the end of the
Cryogenian The Cryogenian (from grc, κρύος, krýos, meaning "cold" and , romanized: , meaning "birth") is a geologic period that lasted from . It forms the second geologic period of the Neoproterozoic Era, preceded by the Tonian Period and followed by ...
geological period The geologic time scale, or geological time scale, (GTS) is a representation of time based on the rock record of Earth. It is a system of chronological dating that uses chronostratigraphy (the process of relating strata to time) and geoc ...
and the more familiar fauna of the Late Ediacaran
Avalon explosion The Avalon explosion, named from the Precambrian fauna of the Avalon Peninsula, is a proposed evolutionary radiation in the history of the Animalia, about 575 million years ago in the Ediacaran Period, with the Avalon explosion being one of th ...
, as well as due to its microfossils' potential utility as biostratigraphical markers. Taken as a whole, the Doushantuo Formation ranges from about 635 Ma (million years ago) at its base to about 551 Ma at its top, with the most fossiliferous layer predating by perhaps five Ma the earliest of the 'classical' Ediacaran faunas from
Mistaken Point Mistaken Point Ecological Reserve is a wilderness area and a UNESCO World Heritage Site located at the southeastern tip of Newfoundland's Avalon Peninsula in the Canadian province of Newfoundland and Labrador. The reserve is home to the namesak ...
on the
Avalon Peninsula The Avalon Peninsula (french: Péninsule d'Avalon) is a large peninsula that makes up the southeast portion of the island of Newfoundland. It is in size. The peninsula is home to 270,348 people, about 52% of Newfoundland's population, according ...
of
Newfoundland Newfoundland and Labrador (; french: Terre-Neuve-et-Labrador; frequently abbreviated as NL) is the easternmost province of Canada, in the country's Atlantic region. The province comprises the island of Newfoundland and the continental region ...
, and recording conditions up to a good forty to fifty million years before the
Cambrian explosion The Cambrian explosion, Cambrian radiation, Cambrian diversification, or the Biological Big Bang refers to an interval of time approximately in the Cambrian Period when practically all major animal phyla started appearing in the fossil recor ...
at the beginning of the
Phanerozoic The Phanerozoic Eon is the current geologic eon in the geologic time scale, and the one during which abundant animal and plant life has existed. It covers 538.8 million years to the present, and it began with the Cambrian Period, when anima ...
.


Sedimentology

The whole sequence sits on an
unconformity An unconformity is a buried erosional or non-depositional surface separating two rock masses or strata of different ages, indicating that sediment deposition was not continuous. In general, the older layer was exposed to erosion for an interval o ...
with the underlying Liantuo formation, which is free of fossils, an unconformity usually being interpreted as a period of erosion. On that unconformity lie tillites of the Nantuo formation - cemented
glacial till image:Geschiebemergel.JPG, Closeup of glacial till. Note that the larger grains (pebbles and gravel) in the till are completely surrounded by the matrix of finer material (silt and sand), and this characteristic, known as ''matrix support'', is d ...
formed of glacial deposits of cobbles and gravel laid down at the end of the
Marinoan glaciation The Marinoan glaciation, sometimes also known as the Varanger glaciation, was a period of worldwide glaciation that lasted from approximately 650 to 632.3 ± 5.9 Ma (million years ago) during the Cryogenian period. This glaciation possibly cover ...
(also known as Varangian glaciation, this is the second and last of a series of very extensive glaciations during a period called the
Cryogenian The Cryogenian (from grc, κρύος, krýos, meaning "cold" and , romanized: , meaning "birth") is a geologic period that lasted from . It forms the second geologic period of the Neoproterozoic Era, preceded by the Tonian Period and followed by ...
-- named because '
Snowball Earth The Snowball Earth hypothesis proposes that, during one or more of Earth's Greenhouse and icehouse Earth, icehouse Climate, climates, the Earth's surface, planet's surface became entirely or nearly entirely Freezing, frozen. It is believed that ...
' conditions at the time). This latest Cryogenian glacial level is tentatively dated ''ca'' 654 (660 ± 5) — 635 Ma (million years ago). The Doushantuo formation itself has three layers representing aquatic sediments that formed as sea levels rose with the melting of worldwide glaciation.
Biomarker In biomedical contexts, a biomarker, or biological marker, is a measurable indicator of some biological state or condition. Biomarkers are often measured and evaluated using blood, urine, or soft tissues to examine normal biological processes, ...
s indicate highly saline conditions, such as might be found in a lagoon, low oxygen levels, and very little sediment that had been washed off land surfaces. The richest finds (the ''Lagerstätte'' itself) lie at the bottom of the middle stratum, with a date about 570 Ma, thus from some time after the great
Gaskiers glaciation The Gaskiers glaciation is a period of widespread glacial deposits (e.g. diamictites) that lasted under 340 thousand years, between 579.63 ± 0.15 and 579.88 ± 0.44 million years ago — i.e. late in the Ediacaran Period — making it the last maj ...
of 85 ± 1 - 582.1 ± 0.4 Ma


Fossils

Doushantuo fossils are all aquatic, microscopic, and preserved to a great degree of detail. The latter two characteristics mean that the structure of the organisms that made them can be studied at the cellular level, and considerable insight has been gained into the embryonic and larval stages of many early creatures. One contentious claim is that many of the fossils show signs of
bilateral symmetry Symmetry in biology refers to the symmetry observed in organisms, including plants, animals, fungi, and bacteria. External symmetry can be easily seen by just looking at an organism. For example, take the face of a human being which has a pla ...
, a common feature in many modern-day animals which is usually assumed to have evolved later, during the Cambrian Explosion. A nearly microscopic fossil animal, ''
Vernanimalcula ''Vernanimalcula guizhouena'' is an acritarch dating from ; it was between 0.1 and 0.2 mm across (roughly the width of one or two human hairs). ''Vernanimalcula'' means "small spring animal", referring to its appearance in the fossil record ...
'' ("springtime micro-animal") was announced in October 2005, with the claim that it was the oldest known bilateral animal. However, the absence of adult forms of almost all animal types in the Doushantuo (there are microscopic adult
sponge Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate through t ...
s and
coral Corals are marine invertebrates within the class Anthozoa of the phylum Cnidaria. They typically form compact colonies of many identical individual polyps. Coral species include the important reef builders that inhabit tropical oceans and sec ...
s) makes these claims difficult to prove: some argue that their lack suggests these finds are not larval and embryonic forms at all; supporters contend that some unidentified process "filtered out" all but the smallest forms from fossilization. An alternative interpretation suggests that it was created by non-biological rock-forming processes. The team that discovered ''Vernanimalcula'' have defended their conclusion that it was an animal, pointing out that they found ten specimens (not illustrated) of the same size and configuration, and stating that non-biological processes would be very unlikely to produce so many specimens that were so alike. The discovery was made when the rich phosphate deposits were being mined, and was first reported in 1998. The finds offer direct evidence that confirms expectations that major evolutionary diversification of animals already had occurred before the onset of the Cambrian period, with its apparent 'explosion' of
metazoan life Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...
-forms and, therefore, that more remote ancestral forms of the phyla recognizable in Cambrian macrofossils must have existed previously. The documented biota now includes phosphatized microfossils of
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
, multicellular
thallophyte Thallophytes (Thallophyta, Thallophyto or Thallobionta) are a polyphyletic group of non-motile organisms traditionally described as "thalloid plants", "relatively simple plants" or "lower plants". They form a division of kingdom Plantae that inc ...
s (seaweeds),
acritarch Acritarchs are organic microfossils, known from approximately 1800 million years ago to the present. The classification is a catch all term used to refer to any organic microfossils that cannot be assigned to other groups. Their diversity refle ...
s,
ciliate The ciliates are a group of alveolates characterized by the presence of hair-like organelles called cilia, which are identical in structure to flagellum, eukaryotic flagella, but are in general shorter and present in much larger numbers, with a ...
s, and cyanophytes, besides adult
sponge Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate through t ...
s and adult
cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that th ...
ns (coelenterates; these may be early forms of
tabulate coral Tabulata, commonly known as tabulate corals, are an order of extinct forms of coral. They are almost always colonial, forming colonies of individual hexagonal cells known as corallites defined by a skeleton of calcite, similar in appearance to ...
s (tetracorallians)). There also seem to be what scientists cautiously report as bilateral animal embryos, termed '' Parapandorina'', and eggs (''
Megasphaera ''Megasphaera'' is a genus of Bacillota bacteria classified within the class Negativicutes. This classification has been reexamined.Yutin N, Galperin MY (2013) A genomic update on Clostridial phylogeny: Gram-negative spore formers and other mi ...
''). Some of the possible animal embryos are in an early stage of cellular division (that was first interpreted as
spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, f ...
s or algal cells), including eggs and embryos which are most probably of sponges or cnidarians, as well as adult sponges and a variety of adult cnidarians. An alternative possibility is that the "embryos" and "eggs" are in fact fossils of giant sulfur
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
resembling ''
Thiomargarita ''Thiomargarita'' is a genus (family Thiotrichaceae) which includes the vacuolate sulfur bacteria species ''Thiomargarita namibiensis'', ''Candidatus Thiomargarita nelsonii'', and ''Ca. Thiomargarita joergensii''. In 2022, scientists working ...
'', a bacterium so large that it is visible to the naked eye. The interpretation would also provide a mechanism for
phosphatic fossilization Phosphatic fossilization has occurred in unusual circumstances to preserve some extremely high-resolution microfossils in which careful preparation can even reveal preserved cellular structures. Such microscopic fossils are only visible under the s ...
through microbially mediated phosphate
precipitation In meteorology, precipitation is any product of the condensation of atmospheric water vapor that falls under gravitational pull from clouds. The main forms of precipitation include drizzle, rain, sleet, snow, ice pellets, graupel and hail. ...
by the bacteria, which has been observed in modern environments. If dark spots in the fossil transpire to be fossilised nuclei - an unlikely claim - this would refute the ''Thiomargarita'' hypothesis. That being said, recent comparisons of the Doushantuo fossils to modern decaying ''Thiomargarita'' and expired sea urchin embryos shows little similarity between the fossils and decaying bacterial cells. Only about one-twentieth of the site's fossils have been excavated. The fossil beds are threatened by increasing intensity of
phosphate mining In chemistry, a phosphate is an anion, salt (chemistry), salt, functional group or ester derived from a phosphoric acids and phosphates, phosphoric acid. It most commonly means orthophosphate, a derivative of phosphoric acid, orthophosphoric a ...
operations in the area. A workshop led in protest by local paleontologists resulted in a temporary halt to the mining in 2017.


Palaeogeography

The formation was laid down on a carbonate shelf, whose rim enclosed a lagoon between tidal flats on the shore, and the deeper ocean. This lagoon was periodically anoxic or
euxinic Euxinia or euxinic conditions occur when water is both anoxic and sulfidic. This means that there is no oxygen (O2) and a raised level of free hydrogen sulfide (H2S). Euxinic bodies of water are frequently strongly stratified, have an oxic, highly ...
(containing
hydrogen sulfide Hydrogen sulfide is a chemical compound with the formula . It is a colorless chalcogen-hydride gas, and is poisonous, corrosive, and flammable, with trace amounts in ambient atmosphere having a characteristic foul odor of rotten eggs. The unde ...
); variations in the chemistry in the lagoon can be detected in isotopic and elemental abundance cycles in the rock and possibly contributed to the fossil preservation.


Geochemistry

The most recent Doushantuo rocks show a sharp decrease in the 13C/12C carbon isotope ratio. Since this change appears to be worldwide but its timing does not match that of any other known major event such as a
mass extinction An extinction event (also known as a mass extinction or biotic crisis) is a widespread and rapid decrease in the biodiversity on Earth. Such an event is identified by a sharp change in the diversity and abundance of multicellular organisms. It ...
, it may represent "possible
feedback Feedback occurs when outputs of a system are routed back as inputs as part of a chain of cause-and-effect that forms a circuit or loop. The system can then be said to ''feed back'' into itself. The notion of cause-and-effect has to be handled ...
relationships between evolutionary innovation and seawater chemistry" in which
metazoans Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...
(multi-celled organisms) removed carbon from the water, which increased the concentration of oxygen, and the increased oxygen level made possible the evolution of new metazoans.


See also

*
Phosphatic fossilization Phosphatic fossilization has occurred in unusual circumstances to preserve some extremely high-resolution microfossils in which careful preparation can even reveal preserved cellular structures. Such microscopic fossils are only visible under the s ...


Footnotes


References

* *Knoll, A. H., 2003. ''Life on a Young Planet''. Princeton Univ. Press. *{{cite journal , doi = 10.1038/35318 , last1 = Xiao , first1 = S. , last2 = Zhang , first2 = Y. , last3 = Knoll , first3 = A. H. , year = 1998 , title = Three-dimensional preservation of algae and animal embryos in a Neoproterozoic phosphorite , journal = Nature , volume = 391 , issue = 6667, pages = 553–558 , bibcode = 1998Natur.391..553X , s2cid = 4350507


External links


Brief overview of Doushantuo formation.
Geologic formations of China Proterozoic Eonothem of Asia Ediacaran Asia Ediacaran geology Phosphorite formations Lagerstätten Fossiliferous stratigraphic units of Asia Paleontology in Guizhou Geography of Guizhou