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Doushantuo Formation

Fossil formation in south-central China


Summary

Fossil formation in south-central China

FieldValue
imageDoushantou Embryo Yinetal2007.jpg
captionA purported Ediacaran embryo contained within an acritarch from the Doushantuo Formation
regionSouth China
periodEdiacaran
ageEdiacaran
~
underliesDengying Formation
prilithologyShale
otherlithologyMudstone, marl, carbonate or phosphate minerals
thicknessUp to 400 m; usually around 200 to 250 m
overliesNantuo Formation
countryChina
namedforDoushantuo, Hubei
namedbyLi Siguang and
year_ts1924

~

The Doushantuo Formation (formerly transcribed as Toushantuo or Toushantou, from ) is a geological formation in western Hubei, eastern Guizhou, southern Shaanxi, central Jiangxi, and other localities in China. It is known for the fossil Lagerstätten in Zigui in Hubei, Xiuning in Anhui, and Weng'an in Guizhou, as one of the oldest beds to contain minutely preserved microfossils, phosphatic fossils that are so characteristic {{cite journal |last1=Awramik |first1=S.M. |last2=McMenamin |first2=D.S. |last3=Yin |first3=C. |last4=Zhao |first4=Z. |last5=Ding |first5=Q.

Sedimentology

The whole sequence sits on an unconformity 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 formed of glacial deposits of cobbles and gravel laid down at the end of the Marinoan glaciation (also known as Varangian glaciation, this is the second and last of a series of very extensive glaciations during a period called the Cryogenian—named because 'Snowball Earth' 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. Biomarkers 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 of [585 ± 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, 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 ("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 sponges and corals) 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.{{cite journal | doi-access = free | name-list-style=amp | doi-access=

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-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, multicellular thallophytes (seaweeds), acritarchs, ciliates, and cyanophytes, besides adult sponges and adult cnidarians (coelenterates; these may be early forms of tabulate corals (tetracorallians)). There also seem to be what scientists cautiously report as bilateral animal embryos, termed Parapandorina, and eggs (Megasphaera). Some of the possible animal embryos are in an early stage of cellular division (that was first interpreted as spores 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 resembling Thiomargarita, a bacterium so large that it is visible to the naked eye. The interpretation would also provide a mechanism for phosphatic fossilization through microbially mediated phosphate precipitation 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 operations in the area. A workshop led in protest by local paleontologists resulted in a temporary halt to the mining in 2017.

Paleobiota

While here they are placed in the Doushantuo Formation, most of the algae are found in the Miaohe biota, which may instead belong to the lower part of the Dengying Formation.

Algae

AlgaeGenusSpeciesHigher taxonNotesImages
AnhuiphytonA. lineatumEukaryotaEssentially identical to specimens from the Lantian Formation
AnomalophytonA. zhangzhongyingiEukaryotaSimilar to Doushantuophyton, but branches less frequently
BaculiphycaB. taeniata, B. brevistipitataEukaryotaSpecies differ in stipe length
BeltanelliformisB. brunsaeCyanobacteriaOriginally interpreted as a polypoid cnidarian[[File:Anthozoa - Nemiana simplex.JPGthumbBeltanelliformis fossil from Ukraine]]
CrassitubusC. costataCyanobacteriaOriginally interpreted as a tubular animal, before being reinterpreted as a cyanobacterium
CobiosC. rubroRhodophytaPreserves evidence of multiple life stages
DoushantuophytonD. lineare, D. rigidulum, D. quyuani, D. cometa, D. laticladusEukaryotaRelatively small and thin
EnteromorphitesE. siniansis, E. magnusEukaryotaOne of the larger "branching algae" from Doushantuo
GemmaphytonG. taoyingensisEukaryotalast1=Wangfirst1=Yelast2=Wangfirst2=Yuelast3=Dufirst3=Weilast4=Wangfirst4=Xunliantitle=New Data of Macrofossils in the Ediacaran Wenghui Biota from Guizhou, South Chinajournal=Acta Geologica Sinicadate=October 2016volume=90issue=5pages=1611–1628url=https://www.researchgate.net/profile/Wei-Du-48/publication/310470096_New_Data_of_Macrofossils_in_the_Ediacaran_Wenghui_Biota_from_Guizhou_South_China/links/59c4b0b2aca272c71bb41abe/New-Data-of-Macrofossils-in-the-Ediacaran-Wenghui-Biota-from-Guizhou-South-China.pdf?_tp=eyJjb250ZXh0Ijp7ImZpcnN0UGFnZSI6InByb2ZpbGUiLCJwYWdlIjoicHVibGljYXRpb24iLCJwcmV2aW91c1BhZ2UiOiJwcm9maWxlIn19}}
GlobusphytonG. lineareEukaryotaFairly unusual morphology suggests it likely crept along the seafloor
GlomulusG. filamentumCyanobacteriaResembles modern cyanobacteria like Microcoleus
GremiphycaG. corymbiataRhodophytaLikely a stem-rhodophyte, resembles coralline algae
GrypaniaG. spiralisEukaryota?One of the most common large Proterozoic fossils[[File:Grypania spiralis.JPGthumbGrypania fossils from the Negaunee Iron Formation in the USA]]
JiuqunaoellaJ. simplicisEukaryotaResembles Grypania, but differs in presence of constrictions
JixianiaJ. lineataEukaryota?Also known from the Mesoproterozoic, probably a filamentous alga
KonglingiphytonK. erecta, K. lateraleRhodophyta?Likely a rhodophyte, a formerly separate genus ("Ramalga") was actually a specimen of this genus overlaid by one of Doushantuophyton
LatiortenuiphytonL. robustaEukaryotaHas large vein-like structures in its thallus
LiulingjitaeniaL. alloplectaEukaryotaUnusually consists of a set of braided filaments
LongifuniculumL. dissolutumEukaryotaSimilar to Liulingjitaenia, but its filaments fray out at each end almost like a braided rope
MaxiphytonM. stipitatumEukaryotalast1=Yefirst1=Qinlast2=Tongfirst2=Jinnanlast3=Anfirst3=Zhihuilast4=Hufirst4=Junlast5=Tianfirst5=Lilast6=Guanfirst6=Kaipinglast7=Xiaofirst7=Shuhaititle=A systematic description of new macrofossil material from the upper Ediacaran Miaohe Member in South Chinajournal=Journal of Systematic Palaeontologydate=1 February 2019volume=17issue=3pages=183–238doi=10.1080/14772019.2017.1404499}}
MegaspirellusM. houiEukaryotaHolotype reinterpreted as a coprolite due to sediment grains
MiaohephytonM. bifurcatumPhaeophyta?Fossils likely represent shed fragments from larger thalli
ParameciaP. incognataCorallinales?Formerly interpreted as a marine lichen
ParatetraphycusP. giganteaBangiophyceae?Resembles early developmental stages of the alga Porphyra
PseudodoushantuophytonP. wenghuiensisEukaryotaBears cluster-like filaments on its branches
QuadratitubusQ. orbigoniatusCyanobacteriaOriginally interpreted as a tubular animal, before being reinterpreted as a cyanobacterium
RamitubusR. increscens, R. decrescensEukaryotalast1=Liufirst1=Pengjulast2=Xiaofirst2=Shuhailast3=Yinfirst3=Chongyulast4=Zhoufirst4=Chuanminglast5=Gaofirst5=Linzhilast6=Tangfirst6=Fengtitle=SYSTEMATIC DESCRIPTION AND PHYLOGENETIC AFFINITY OF TUBULAR MICROFOSSILS FROM THE EDIACARAN DOUSHANTUO FORMATION AT WENG'AN, SOUTH CHINAjournal=Palaeontologydate=March 2008volume=51issue=2pages=339–366doi=10.1111/j.1475-4983.2008.00762.x}} then found to be an alga similar to Epiphyton
SinocylindraS. linearis, S. yunnanensisEukaryotaResembles the cyanobacteria Siphonophycus[[File:Sinocylindra yunnanensis, Chengjiang biota 2.jpgthumbS. yunnanensis fossil from the Chengjiang biota]]
SinocyclocyclicusS. guizhouensisCyanobacterialast1=Sunfirst1=Wei-Chenlast2=Yinfirst2=Zong-Junlast3=Donoghuefirst3=Philiplast4=Liufirst4=Peng-Julast5=Shangfirst5=Xiao-Donglast6=Zhufirst6=Mao-Yantitle=Tubular microfossils from the Ediacaran Weng'an Biota (Doushantuo Formation, South China) are not early animalsjournal=Palaeoworlddate=December 2019volume=28issue=4pages=469–477doi=10.1016/j.palwor.2019.04.004hdl=1983/2fa05771-9d96-4663-8438-29d52f2cc197hdl-access=free }}
SiphonophycusS. solidum, S. septatum, S. kestron, S. typicum, S. robustumCyanobacteriaRelatively common cyanobacteria
ThallophycaT. corrugataRhodophytaResembles the Solenoporaceae
ThallophycoidesT. phloeatusEukaryotaMay be a rhodophyte?
TongrenphytonT. kommaEukaryotaHas several filaments along its thallus, showing a clear eukaryotic affinity
VendotaeniaV. spVendotaenidResembles specimens from the lower Cambrian[[File:Vendotaenid indet.jpgthumbIndeterminate vendotaenid fossil]]
WenganiaW. globosa, W. exquisita, W. minutaEukaryotaResembles various algae in its parenchymatous structure, especially green algae

Microorganisms

MicroorganismsGenusSpeciesHigher taxonNotesImages
AggregatosphaeraA. miaoheensisEukaryota?last1=Xiaofirst1=Shuhailast2=Yuanfirst2=Xunlailast3=Steinerfirst3=Michaellast4=Knollfirst4=Andrew H.title=Macroscopic Carbonaceous Compressions in a Terminal Proterozoic Shale: A Systematic Reassessment of the Miaohe Biota, South Chinajournal=Journal of Paleontologydate=2002volume=76issue=2pages=347–376url=https://www.jstor.org/stable/pdf/1307146.pdfissn=0022-3360}}
AnnularidensA. inconditusAcritarchlast1=Yefirst1=Qinlast2=Lifirst2=Jiaqilast3=Tongfirst3=Jinnanlast4=Anfirst4=Zhihuilast5=Hufirst5=Junlast6=Xiaofirst6=Shuhaititle=A microfossil assemblage from the Ediacaran Doushantuo Formation in the Shennongjia area (Hubei Province, South China): Filling critical paleoenvironmental and biostratigraphic gapsjournal=Precambrian Researchdate=August 2022volume=377article-number=106691doi=10.1016/j.precamres.2022.106691}} giving it a gearwheel-like appearance in cross-section
ApodastoidesA. basileusAcritarchBears processes with plugs at their base
AppendisphaeraA. anguina, A. clava, A. grandis, A. fragilis, A. longispina, A. setosa, A. tabifica, A. tenuis, A. helicaea, A. brevispina, A. hemisphaericaAcritarchBears numerous long, flexible processes
ArchaeophycusA. yunnanensisEukaryota?Either a cyanobacteria or the early developmental stage of an alga
AsterocapsoidesA. sinicusAcritarchOriginal holotype was damaged by the surrounding rock breaking
BacatisphaeraB. baokangensisAcritarchlast1=Chuanmingfirst1=Zhoulast2=Brasierfirst2=M. D.last3=Yaosongfirst3=Xuetitle=Three‐Dimensional Phosphatic Preservation Of Giant Acritarchs From The Terminal Proterozoic Doushantuo Formation In Guizhou And Hubei Provinces, South Chinajournal=Palaeontologydate=November 2001volume=44issue=6pages=1157–1178doi=10.1111/1475-4983.00219}}
BaltisphaeridiumB. rigidumAcritarchlast1=Xunlaifirst1=Yuanlast2=Hofmannfirst2=H.J.title=New microfossils from the neoproterozoic (Sinian) Doushantuo Formation, Wengan, Guizhou Province, southwestern Chinajournal=Alcheringa: An Australasian Journal of Palaeontologydate=January 1998volume=22issue=3pages=189–222doi=10.1080/03115519808619200}}
BispinosphaeraB. vacuaAcritarchBears two separate types of process; one is conical and hollow, the other is solid and hair-like
BotominellaB. lineataEukaryota?Known from small trichome-like structures
CastaneasphaeraC. speciosaAcritarchResembles Phanerozoic "mazuelloids"
CavaspiraC. acuminata, C. basiconicaAcritarchBears short, conical processes
CerionoporaC. ordinataAcritarchMay be a resting cyst of a multicellular alga
ComasphaeridiumC. magnumAcritarchBears long hair-like processes
CrassimembranaC. crispans, C. multitunicaAcritarchlast1=Ouyangfirst1=Qinglast2=Zhoufirst2=Chuanminglast3=Xiaofirst3=Shuhailast4=Guanfirst4=Chengguolast5=Chenfirst5=Zhelast6=Yuanfirst6=Xunlailast7=Sunfirst7=Yunpengtitle=Distribution of Ediacaran acanthomorphic acritarchs in the lower Doushantuo Formation of the Yangtze Gorges area, South China: Evolutionary and stratigraphic implicationsjournal=Precambrian Researchdate=February 2021volume=353article-number=106005doi=10.1016/j.precamres.2020.106005}}
CyanonemaC. attenuatumCyanobacteriaSimilar to forms from the Bitter Springs formation
CymatiosphaeroidesC. forabilatus, C. yiniiAcritarchSimilar to Membranosphaera, bears cylindrical processes with pointed tips
DicrospinasphaeraD. zhangiiAcritarchBears thin, branching processes
DistosphaeraD. jinguadunensis, D. speciosaAcritarchBears two layers of cylindrical processes
DoushantuonemaD. peatiiCyanobacteriaEarliest reported cyanobacteria from the formation
DuospinosphaeraD. shennongjiaensis, D. biformisAcritarchBears two layers of processes; an inner cylindrical one likened to hanging icicles and a larger conical one on the outer wall
EchinosphaeridiumE. maximumAcritarchFormerly placed in Ericiasphaera due to a misinterpretation of the spines as solid
EotylotopallaE. apophysa, E. dactylos, E. delicataAcritarchFormerly placed within Timanisphaera
EriciasphaeraE. fibrila, E. magna, E. rigida?, E. densispina, E. sparsaAcritarchlast1=Dunthornfirst1=Micahlast2=Lippsfirst2=Jere H.last3=Stoeckfirst3=Thorstentitle=Reassessment of the Putative Ciliate Fossils Eotintinnopsis, Wujiangella, and Yonyangella from the Neoproterozoic Doushantuo Formation in Chinajournal=Acta Protozoologicadate=2010volume=49issue=2pages=139–144url=https://ejournals.eu/en/journal/acta-protozoologica/article/reassessment-of-the-putative-ciliate-fossils-eotintinnopsis-wujiangella-and-yonyangella-from-the-neoproterozoic-doushantuo-formation-in-chinalanguage=en}}
GoniosphaeridiumG. acuminatum, G. conoideum, G. cratumAcritarchSimilar to Baltisphaeridium
GranitunicaG. mcfaddenaeAcritarchDiffers from other sphaeromorph acritarchs in its granular wall
HocosphaeridiumH. dilatatum, H. scaberfaciumAcritarchBears numerous hooked processes
KnollisphaeridiumK. maximum, K. coniformum, K. denticulatum, K. longilatum, K. obtusum, K. parvumAcritarchBears conical processes with a long filament-like tip
LeiosphaeridiaL. tenuissima, L. crassa, L. minutissimaAcritarchIncredibly abundant
MegasphaeraM. inornataProtistaThought represent an animal egg, then revealed to likely be an encysting protist
MeghystricosphaeridiumM. chadianensis, M?. densum, M. gracilentum, M. perfectum, M. magnificum, M. wenganensisAcritarchSomewhat unclear which species belong to this genus
MengeosphaeraM. angusta?, M. chadianensis, M. constricta, M. gracilis, M. mamma, M. minima, M. stegosauriformis, M. bellula, M. cuspidata, M. grandispina, M. latibasis, M. spicata, M. spinula, M. triangula, M. uniformis, M. membraniferaAcritarchlast1=Liufirst1=Pengjulast2=Xiaofirst2=Shuhailast3=Yinfirst3=Chongyulast4=Chenfirst4=Shouminglast5=Zhoufirst5=Chuanminglast6=Lifirst6=Mengtitle=Ediacaran Acanthomorphic Acritarchs and Other Microfossils from Chert Nodules of the Upper Doushantuo Formation in the Yangtze Gorges Area, South Chinajournal=Journal of Paleontologydate=January 2014volume=88issue=S72pages=1–139doi=10.1666/13-009}}
ObruchevellaO. minorCyanobacteriaRelatively rare
OscillatoriopsisO. amadeus, O. longa, O. obtusa, O. majusculaCyanobacteriaCommon cyanobacterial genus
OsculosphaeraO. arcelliformis, O. hyalina, O. membraniferaAcritarchResembles arcellinids
PapillomembranaP. comptaAcritarchFormerly placed within Dasycladales
PustulisphaeraP. membranaceaAcritarchBears three wall layers, with the middle one bearing pimple-like processes
SalomeS. hubeiensisCyanobacteriaA fossil identified as a "microburrow" was placed in this genus before being reidentified as an oblique section of a tubular fossil
SarcinophycusS. radiatus, S. papilloformisEukaryota?Only known from Doushantuo, consists of bundles of sarcinoid cell packets
SchizofusaS. zangwenlongiiAcritarchSimilar to Leiosphaeridia
SinosphaeraS. asteriformis, S. rupinaAcritarchBears small conical spines and a few larger ones
SymphysosphaeraS. basimembranaAcritarchSimilar to Eosphaera, but larger
TanariumT. acus, T. conoideum, T. elegans, T. longitubulare, T. minimum, T. obesum, T. pilosiusculum, T. pycnacanthum, T. variumAcritarchSpine shapes vary from small and triangular to long and needle-like
TianzhushaniaT. spinosaAcritarchBears a many-layered wall with spines penetrating through it
UrasphaeraU. fungiformis, U. nuptaAcritarchBears mushroom-shaped spines, hence the specific name fungiformis
VariomargosphaeridiumV. floridumAcritarchBears processes which branch at their tips
VulcanosphaeraV. phacelosaAcritarchBears bundled hair-like processes
WeissiellaW. grandistellaAcritarchDoushantuo fossils resemble Russian specimens, but are smaller
XenosphaeraX. liantuoensisAcritarchBears extremely thin processes, which have a tendency to not preserve
YushengiaY. ramispinaAcritarchResembles Weissiella, but has longer spines

Miscellaneous taxa

Miscellaneous taxaGenusSpeciesHigher taxonNotesImages
CucullusC. fraudulentusincertae sedisOriginally interpreted as a sponge, then reinterpreted as a microbialite
ParagraptobrancaP. curvusincertae sedisShares features with both graptolites and macroalgae
CalyptrinaC. striataAnnelida?Almost certainly a metazoan[[File:Calyptrina striata sketch.jpgthumbCalyptrina reconstruction]]
EoandromedaE. octobrachiataCtenophora?Originally interpreted as the adult stage of various embryos, then as a pelagic discoid ctenophore, then an umbrella-shaped demersal form[[File:Eoandromeda.jpegthumbleftArtist restoration of the "discoid ctenophore" hypothesis]]
EocyathispongiaE. qianiaPoriferaEarliest likely sponge fossil[[File:Eocyathispongia_qiani.jpgthumbReconstruction of Eocyathispongia]]
LinbotulitaeniaMetazoaLikely a trace fossil of a wriggling mucus-covered animal, possibly from Wenghuiia
ProtoconitesP. minorCnidaria?Likely a cnidarian-grade organism similar to Cambrorhytium[[File:Protoconites 3D Reconstruction.pngthumbReconstruction of Protoconites]]
SinospongiaS. chenjunyuani, S. typicaPorifera?May also be a vermiform alga like the Huainan biota
Trilobozoa indet.UnapplicableMetazoaKnown from an undescribed triradial fossil
VernanimalculaV. guizhouensisBilateria?last1=Petryshynfirst1=Victoria A.last2=Bottjerfirst2=David J.last3=Chenfirst3=Jun-Yuanlast4=Gaofirst4=Fengtitle=Petrographic analysis of new specimens of the putative microfossil Vernanimalcula guizhouena (Doushantuo Formation, South China)journal=Precambrian Researchdate=February 2013volume=225pages=58–66doi=10.1016/j.precamres.2011.08.003}}
WenghuiiaW. jiangkouensisAnnelida?Putatively the earliest known annelid (and also the earliest known bilaterian), even though it already seems very derived

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 (containing hydrogen sulfide); 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, it may represent "possible feedback relationships between evolutionary innovation and seawater chemistry" in which metazoans (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.{{ cite journal | doi-access=free

Footnotes

References

  • Knoll, A. H., 2003. Life on a Young Planet. Princeton Univ. Press.

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