Dickinsonia: Difference between revisions

Dickinsonia; Temporal range: Late Ediacaran, 567–550 Ma PreꞒ Ꞓ O S D C P T J K Pg N
	Cast of Dickinsonia costata from Australia
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	†Proarticulata
Class:	†Dipleurozoa
Family:	†Dickinsoniidae
Genus:	†Dickinsonia; Sprigg, 1947
Type species
	†Dickinsonia costata; Sprigg, 1947
Species
	D. costata (Sprigg, 1947); D. menneri (Keller, 1976); D. tenuis (Glaessner & Wade, 1966);
Synonyms
	Genus synonymy Chondroplon? (Wade, 1971) ; Papilionata Sprigg, 1947 ; Vendomia Keller, 1976 ; D. costata synonymy Papilionata eyrei (Sprigg, 1947) ; D. minima (Sprigg, 1949 ; D. elongata (Glaessner & Wade, 1966) ; D. spriggi (Harrington & Moore, 1955) ; D. tenuis synonymy D. brachina (Wade, 1972) ; D. lissa (Wade, 1972) ; D. rex (Jenkins, 1992) ;

Browse history interactively

← Previous edit Next edit →

Content deleted Content added

VisualWikitext

Inline

Revision as of 06:40, 24 April 2024

Dickinsonia is a genus of extinct organism, most likely an animal, that lived during the late Ediacaran period in what is now Australia, China, Russia, and Ukraine. It is one of the best known members of the Ediacaran biota. The individual Dickinsonia typically resembles a bilaterally symmetrical ribbed oval. Its affinities are presently unknown; its mode of growth has been considered consistent with a stem-group bilaterian affinity,^[3] though various other affinities have been proposed.^[4]^[5]^[6] It lived during the late Ediacaran (final part of Precambrian).^[7] The discovery of cholesterol molecules in fossils of Dickinsonia lends support to the idea that Dickinsonia was an animal,^[8] though these results have been questioned.^[9]

Description

Dickinsonia fossils are known only in the form of imprints and casts in sandstone beds. The specimens found range from a few millimetres to about 1.4 metres (4 ft 7 in) in length, and from a fraction of a millimetre to a few millimetres thick.^[10] They are nearly bilaterally symmetric, segmented, round or oval in outline, slightly expanded to one end (i.e. egg-shaped outline). The rib-like segments are radially inclined towards the wide and narrow ends, and the width and length of the segments increases towards the wide end of the fossil.^[2]^[11] The body is divided into two by a midline ridge or groove,^[2]^[11]^[12] except for a single unpaired segment at one end, dubbed the "anterior most unit" suggested to represent the front of the organism.^[12] Is it disputed whether the segments are offset from each other following glide reflection, and are thus isomers,^[2]^[11]^[13]^[14] or whether the segments are symmetric across the midline, and thus follow true bilateral symmetry, as the specimens displaying the offset may be the result of taphonomic distortion.^[12]^[15] The number of segments/isomer pairs varies from 12 in smaller individuals to 74 in the largest Australian specimens.^[15]

The body of Dickinsonia is suggested to have been sack-like, with the outer layer being made of a resistant but unmineralised material.^[14] Some specimens from Russia show the presence of branched internal structures.^[16]^[14] Some authors have suggested that the underside of the body bore cilia, as well as infolded pockets.^[14]

Dickinsonia is suggested to have grown by adding a new pair of segments/isomers at the end opposite the unpaired "anterior most unit".^[12]^[17] Dickinsonia probably exhibited indeterminate growth (having no maximum size), though it is suggested that the addition of new segments slowed down later in growth.^[18] Deformed specimens from Russia indicate that individuals of Dickinsonia could regenerate after being damaged.^[17]

Ontogeny of Dickinsonia costata following the glide reflection interpretation
Growth of D. costata under bilateral symmetry interpretation
Diagram of various Dickinsonia species
Diagram of various Dickinsonia species (cont)
Diagram of branched internal structures observed in Russian specimens

Ecology

Dickinsonia is suggested to have been a mobile marine organism that lived on the seafloor and fed by consuming microbial mats growing on the seabed using structures present on its underside. Dickinsonia-shaped trace fossils, presumed to represent feeding impressions, sometimes found in chains demonstrating this behaviour have been observed.^[14] These trace fossils have been assigned to the genus Epibaion.^[13]^[19]^[20] A 2022 study suggested that Dickinsonia temporarily adhered itself to the seafloor by the use of mucus, which may have been an adaptation to living in very shallow water environments.^[21]

Discovery

The first species and specimens of this fossil organism were first discovered in the Ediacara Member of the Rawnsley Quartzite, Flinders Ranges in South Australia. Reg Sprigg, the original discoverer of the Ediacaran biota in Australia,^[22] described Dickinsonia, naming it after Ben Dickinson, then Director of Mines for South Australia, and head of the government department that employed Sprigg.^[23] Additional specimens of Dickinsonia are also known from the Mogilev Formation in the Dniester River Basin of Podolia, Ukraine,^[24] the Lyamtsa, Verkhovka, Zimnegory and Yorga Formations in the White Sea area of the Arkhangelsk Region, Chernokamen Formation of the Central Urals, Russia,^[10] (these deposits have been dated to 567–550 Myr.^[25]^[26]^[27]), the Dengying Formation in the Yangtze Gorges area, South China. (ca. 551–543 Ma).^[28]

Taphonomy

As a rule, Dickinsonia fossils are preserved as negative impressions ("death masks") on the bases of sandstone beds. Such fossils are imprints of the upper sides of the benthic organisms that have been buried under the sand.^[29]^[30] The imprints formed as a result of cementation of the sand before complete decomposition of the body. The mechanism of cementation is not quite clear; among many possibilities, the process could have arisen from conditions which gave rise to pyrite "death masks"^[30] on the decaying body, or perhaps it was due to the carbonate cementation of the sand.^[31] The imprints of the bodies of organisms are often strongly compressed, distorted, and sometimes partly extend into the overlying rock. These deformations appear to show attempts by the organisms to escape from the falling sediment.^[13]^[19]^[32]

Rarely, Dickinsonia have been preserved as a cast in massive sandstone lenses, where it occurs together with Pteridinium, Rangea and some others.^[33]^[34]^[35]^[36] These specimens are products of events where organisms were first stripped from the sea-floor, transported and deposited within sand flow.^[33]^[36] In such cases, stretched and ripped Dickinsonia occur. The first such specimen was described as a separate genus and species, Chondroplon bilobatum^[37] and later re-identified as Dickinsonia.

Taxonomy

Species

Since 1947, a total of nine species have been described, of which three are currently considered valid:^[38]

Species	Authority	Location	Status	Notes	Refs
Dickinsonia brachina	Wade (1972)	Australia	invalid	synonym of D. tenuis	^[39]
Dickinsonia costata	Sprigg (1947a)	Australia, Russia, and Ukraine	valid	^[A]	^[40]
Dickinsonia elongata	Glaessner & Wade (1966)	Australia	invalid	synonym of D. costata	^[41]
Dickinsonia lissa	Wade (1972)	Australia	invalid	synonym of D. tenuis	^[39]
Dickinsonia menneri	Keller (1976)	Russia	valid	^[B]^[C]	^[42]
Dickinsonia minima	Sprigg (1949)	Australia	invalid	synonym of D. costata	^[43]
Dickinsonia rex	Jenkins (1992)	Australia	invalid	synonym of D. tenuis	^[44]
Dickinsonia spriggi	Harrington & Moore (1955)	Australia	invalid	synonym of D. costata	^[45]
Dickinsonia tenuis	Glaessner & Wade (1966)	Australia and Russia	valid	^[D]	^[41]

^ Unlike other species, D. costata has comparatively rounded body and fewer, wider segments / isomers.
^ Dickinsonia menneri originally was identified as Vendomia but re-classified as Dickinsonia by Ivantsov (2007)^[2]
^ D. menneri is a small organism up to 8 mm in length, and strongly resembles juvenile specimens of D. costata with its small number of isomers and well-marked head. D. menneri differs from juvenile D. costata by its slightly more elongated form.
^ Dickinsonia tenuis strongly resembles D. costata, but differs from it by more narrow and numerous segments, sparingly lengthened oval form of the body.

A claimed specimen of Dickinsonia from India was later determined to be the remains of a beehive.^[46]

External relationships

Dickinsonia is classified as part of the group Proarticulata or Dickinsoniomorpha.^[14] Proarticulata includes a number of morphologically similar organisms, such Spriggina, Yorgia, Andiva and Cephalonega, which share the same segmented articulation.^[47] The affinities of Proarticulata to other organisms, including to other members of the Ediacaran biota, like rangeomorphs, have long been contentious.^[8] It has been historically proposed that most Ediacaran organisms were closely related to each other, as part of the grouping "Vendobionta",^[5] though recent authors argue that this grouping as a whole is likely to be polyphyletic.^[8] Gregory Retallack has proposed that the fossils of Dickinsonia and other Ediacaran biota represent lichens that grew in a terrestrial environment,^[48] but this has been broadly rejected by other authors, who argue that a marine environment of deposition better fits available evidence.^[49]^[8]^[50] Other proposal have included giant protists, as proposed by Adolf Seilacher.^[51] Most modern research suggest that Dickinsonia and other proarticulatans are likely to be animals, possibly belonging to Eumetazoa.^[18]^[12]^[14] A chemical study of Russian specimens found that they were enriched with cholesterol, which is only produced by animals, supporting an animal affinity,^[8] though these results have been questioned by other authors, who consider the association between the cholesterol molecules and the Dickinsonia fossils to not be definitive.^[9] Within Animalia, a number of affinities have been proposed, including as stem-eumetazoans forming a clade with rangeomorphs,^[52] to Placozoa,^[53] and to Cnidaria.^[54] A number of researchers have proposed close affinities to Bilateria, based on the bilateral or nearly bilateral organisation of proarticulatans,^[14]^[3] though proarticulatans are not likely to be a member of the bilaterian crown group.^[12]

References

^ Hofmann, Hans J. (1988). "An alternative interpretation of the Ediacaran (Precambrian) chondrophore Chondroplon (Wade)". Alcheringa. 12 (4): 315–318. doi:10.1080/03115518808619130.
^ ^a ^b ^c ^d ^e Ivantsov, A.Y. (2007a). "Small Vendian transversely articulated fossils". Paleontological Journal. 41 (2): 113–122. doi:10.1134/S0031030107020013. S2CID 86636748.
^ ^a ^b Gold, D.A.; Runnegar, B.; Gehling, J.G.; Jacobs, D.K. (2015). "Ancestral state reconstruction of ontogeny supports a bilaterian affinity for Dickinsonia". Evolution & Development. 17 (6): 315–397. doi:10.1111/ede.12168. PMID 26492825. S2CID 26099557.
^ Pflug (1973). "Zur fauna der Nama-Schichten in Südwest-Afrika. IV. Mikroscopische anatomie der petalo-organisme" [On the fauna of the Nama layers in southwest Africa. [part] IV. Microscopic anatomy of petaloorganisms]. Palaeontographica (B144): 166–202.
^ ^a ^b Seilacher, Adolf (1992). "Vendobionta and Psammocorallia: Lost constructions of Precambrian evolution". Journal of the Geological Society, London. 149 (4): 607–613. Bibcode:1992JGSoc.149..607S. doi:10.1144/gsjgs.149.4.0607. S2CID 128681462. Retrieved 21 June 2007.
^ McMenamin, M. (1998). The Garden of Ediacara. New York: Columbia University Press. ISBN 978-0-231-10559-0. OCLC 228271905.
^ Dinosaurs: A visual encyclopedia. New York, NY: DK Publishing. 3 April 2018. ISBN 9781465469489.
^ ^a ^b ^c ^d ^e Bobrovskiy, Ilya; Hope, Janet M.; Ivantsov, Andrey; Nettersheim, Benjamin J.; Hallmann, Christian; Brocks, Jochen J. (20 September 2018). "Ancient steroids establish the Ediacaran fossil Dickinsonia as one of the earliest animals". Science. 361 (6408): 1246–1249. Bibcode:2018Sci...361.1246B. doi:10.1126/science.aat7228. hdl:1885/230014. PMID 30237355.
^ ^a ^b Love, G.D.; Zumberge, J.A. (2021). "Emerging patterns in Proterozoic lipid biomarker records". Cambridge Elements. 361 (6408). doi:10.1017/9781108847117. ISBN 9781108847117.
^ ^a ^b Fedonkin, M.A.; Gehling, J.G.; Grey, K.; Narbonne, G.M.; Vickers-Rich, P. (2007). The Rise of Animals: Evolution and diversification of the kingdom Animalia. Johns Hopkins University Press. p. 326. ISBN 978-0-8018-8679-9.
^ ^a ^b ^c Ivantsov, A. Yu (2012). "Becoming metamery and bilateral symmetry in Metazoa: Way of Proarticulata". Morphogenesis in the Individual and Historical Development: Symmetry and asymmetry: 16–17.
^ ^a ^b ^c ^d ^e ^f Evans, Scott D.; Droser, Mary L.; Gehling, James G. (17 May 2017). Hejnol, Andreas (ed.). "Highly regulated growth and development of the Ediacara macrofossil Dickinsonia costata". PLoS One. 12 (5): e0176874. doi:10.1371/journal.pone.0176874. ISSN 1932-6203. PMC 5435172. PMID 28520741.
^ ^a ^b ^c Ivantsov, A.Y. (2011). "Feeding traces of Proarticulata — the Vendian metazoa". Paleontological Journal. 45 (3): 237–248. doi:10.1134/S0031030111030063. S2CID 128741869.
^ ^a ^b ^c ^d ^e ^f ^g ^h Ivantsov, Andrey Yu; Zakrevskaya, Maria (23 February 2023). "Body plan of Dickinsonia , the oldest mobile animals". Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 114 (1–2): 95–108. doi:10.1017/S175569102300004X. ISSN 1755-6910.
^ ^a ^b Dunn, Frances S.; Liu, Alexander G.; Donoghue, Philip C.J. (May 2018). "Ediacaran developmental biology". Biological Reviews. 93 (2): 914–932. doi:10.1111/brv.12379. ISSN 1464-7931. PMC 5947158. PMID 29105292.
^ Ivantsov, A.Y. (2004). "New Proarticulata from the Vendian of the Arkhangel'sk region" (PDF). Paleontological Journal. 38 (3): 247–253. Archived from the original (PDF) on 27 September 2007. Retrieved 27 September 2007.
^ ^a ^b Ivantsov, Andrey; Zakrevskaya, Maria; Nagovitsyn, Aleksey; Krasnova, Anna; Bobrovskiy, Ilya; Luzhnaya (Serezhnikova), Ekaterina (November 2020). "Intravital damage to the body of Dickinsonia (Metazoa of the late Ediacaran)". Journal of Paleontology. 94 (6): 1019–1033. doi:10.1017/jpa.2020.65. ISSN 0022-3360.
^ ^a ^b Evans, Scott D.; Hunt, Gene; Gehling, James G.; Sperling, Erik A.; Droser, Mary L. (January 2023). Rahman, Imran (ed.). "Species of Dickinsonia Sprigg from the Ediacaran of South Australia". Palaeontology. 66 (1). doi:10.1111/pala.12635. ISSN 0031-0239.
^ ^a ^b Ivantsov, A.Y. (2013). "Trace Fossils of Precambrian Metazoans "Vendobionta" and "Mollusks"". Stratigraphy and Geological Correlation. 21 (3): 252–264. Bibcode:2013SGC....21..252I. doi:10.1134/S0869593813030039. S2CID 128638405.
^ Ivantsov, A.Y.; Malakhovskaya, Y.E. (2002). "Giant Traces of Vendian Animals" (PDF). Doklady Earth Sciences. 385 (6): 618–622. Archived from the original (PDF) on 4 July 2007. Retrieved 24 February 2008.
^ Ivantsov, Andrey; Zakrevskaya, Maria (July 2022). "Dickinsonia : mobile and adhered". Geological Magazine. 159 (7): 1118–1133. doi:10.1017/S0016756821000194. ISSN 0016-7568.
^ Sprigg, Reg C. (1947). "Early Cambrian (?) jellyfishes from the Flinders Ranges, South Australia" (PDF). Trans. R. Soc. S. Aust. 71: 212–24. Archived from the original (PDF) on 29 September 2007.
^ Eig, Karsten. "EDIACARANS: THE FOSSILS THAT SHOULD NOT BE THERE". Adventures in geology - Karsten Eig.
^ Fedonkin, M. A. (1983). "Non-skeletal fauna of Podoloia, Dniester River valley". In Velikanov, V. A.; Assejeva, E. A.; Fedonkin, M. A. (eds.). The Vendian of the Ukraine (in Russian). Kiev: Naukova Dumka. pp. 128–139.
^ Grazhdankin, Dima (2004). "Patterns of distribution in the Ediacaran biotas: facies versus biogeography and evolution" (PDF). Paleobiology. 30 (2): 203–221. doi:10.1666/0094-8373(2004)030<0203:PODITE>2.0.CO;2. S2CID 129376371.
^ Maslov, A.V.; Podkovyrov, V.N.; Grazhdankin, D.V.; Kolesnikov, A.V. (2018). "Upper Vendian in the east, northeast and north of East European Platform: Depositional processes and biotic evolution". Litosfera. 18 (4): 520–542. doi:10.24930/1681-9004-2018-18-4-520-542.
^ Kolesnikov, A.V.; Liu, A.G.; Danelian, T.; Grazhdankin, D.V. (2018). "A reassessment of the problematic Ediacaran genus Orbisiana Sokolov 1976". Precambrian Research. 316: 197–205. Bibcode:2018PreR..316..197K. doi:10.1016/j.precamres.2018.08.011. S2CID 134213721.
^ Wang, Xiao-Peng; Chen, Zhe; Pang, Ke; Zhou, Chuan-Ming; Xiao, Shuhai; Wan, Bin; Yuan, Xun-Lai (2021). "Dickinsonia from the Ediacaran Dengying Formation in the Yangtze Gorges area, South China". Palaeoworld. 30 (4): 602–609. doi:10.1016/j.palwor.2021.01.002. S2CID 234254399.
^ Ivantsov, A.Y. (2009). "A new reconstruction of Kimberella, a problematic Vendian Metazoan". Paleontological Journal. 43 (6): 601–611. doi:10.1134/S003103010906001X. S2CID 85676210.
^ ^a ^b Gehling, J.G. (1999). "Microbial mats in terminal Proterozoic siliciclastics; Ediacaran death masks". PALAIOS. 14 (1): 40–57. Bibcode:1999Palai..14...40G. doi:10.2307/3515360. JSTOR 3515360.
^ Serezhnikova Z, E.A. (2011). "Microbial Binding as a Probable Cause of Taphonomic Variability of Vendian Fossils: Carbonate Casting?". Advances in Stromatolite Geobiology. Lecture Notes in Earth Sciences. Vol. 131. pp. 525–535. doi:10.1007/978-3-642-10415-2_31. ISBN 978-3-642-10414-5.
^ Runnegar, Bruce (1982). "Oxygen requirements, biology and phylogenetic significance of the late Precambrian worm Dickinsonia, and the evolution of the burrowing habit". Alcheringa: An Australasian Journal of Palaeontology. 6 (3): 223–239. doi:10.1080/03115518208565415.
^ ^a ^b Grazhdankin, Dima (2004). "Patterns of distribution in the Ediacaran biotas: facies versus biogeography and evolution" (PDF). Paleobiology. 30 (2): 203–221. doi:10.1666/0094-8373(2004)030<0203:PODITE>2.0.CO;2. S2CID 129376371.
^ Keller, B.M.; Fedonkin, M.A. (1976). "New records of fossils in the Valdaian group of the precambrian on the Syuz'ma River" (PDF). Izvestiya Akademii Nauk SSR. Seriya Geologicheskaya (in Russian). 3: 38–44. Archived from the original (PDF) on 27 September 2007.
^ Keller, B. M.; Fedonkin, M. A. (1977). "New organic fossil finds in the Precambrian Valday series along the Syuz'ma River". International Geology Review. 19 (8): 924–930. Bibcode:1977IGRv...19..924K. doi:10.1080/00206817709471091.
^ ^a ^b Gehling, J.G.; Droser, M.L. (2013). "How well do fossil assemblages of the Ediacara Biota tell time?". Geology. 41 (4): 447–450. Bibcode:2013Geo....41..447G. doi:10.1130/G33881.1.
^ Wade, M. (1971). "Bilateral Precambrian Chondrophores from the Ediacara Fauna, South Australia". Proceedings of the Royal Society of Victoria. 84 (1): 183–188.
^ Evans, S.D.; Hunt, G.; Gehling, J.G.; Sperling, E. A.; Droser, M.L. (2023). "Species of Dickinsonia sprigg from the Ediacaran of South Australia". Palaeontology. 66 (1–21): e12635. doi:10.1111/pala.12635.
^ ^a ^b Wade, M. (1972). "Dickinsonia: Polychaete worms from the late Precambrian Ediacara fauna, South Australia". Mem. Queensl. Mus. 16 (2): 171–190.
^ Sprigg, Reg C. (1947a). "Early Cambrian (?) jellyfishes from the Flinders Ranges, South Australia" (PDF). Trans. R. Soc. S. Aust. 71: 212–24. Archived from the original (PDF) on 29 September 2007.
^ ^a ^b Glaessner, M.F.; Wade, M. (1966). "The late Precambrian fossils from Ediacara, South Australia" (PDF). Palaeontology. 9 (4): 599.
^ Keller, B.M.; Fedonkin, M.A. (1976). "New records of fossils in the Valdaian group of the precambrian on the Syuz'ma River" (PDF). Izvestiya Akademii Nauk SSR. Seriya Geologicheskaya (in Russian). 3: 38–44. Archived from the original (PDF) on 27 September 2007.
^ Sprigg, R.C. (1949). "Early Cambrian "jellyfishes" of Ediacara, South Australia, and Mount John, Kimberley District, Western Australia" (PDF). Transactions of the Royal Society of South Australia. 73: 72–99.{{cite journal}}: CS1 maint: url-status (link)^{[permanent dead link]}
^ Jenkins, R.J.F. (1992). "Functional and ecological aspects of Ediacarian assemblages". In Lipps, J.; Signor, P.W. (eds.). Origin and Early Evolution of the Metazoa. New York, NY: Springer. pp. 131–176. ISBN 978-0-306-44067-0. OCLC 231467647.
^ Harrington, N.J.; Moore, R.C. (1955). "Kansas Pennsylvanian and other jellyfishes". Bull. Kansas Geol. Surv. 114 (5): 153–163.
^ Pandey, S.K.; Ahmad, Shamim; Sharma, Mukund (9 March 2023). "Dickinsonia tenuis reported by Retallack et al. 2021 is not a fossil, instead an impression of an extant 'fallen beehive'". Journal of the Geological Society of India. 99 (3): 311–316. doi:10.1007/s12594-023-2312-2. ISSN 0974-6889.
^ Ivantsov, A.Y.; Fedonkin, M.A.; Nagovitsyn, A.L.; M.A. Zakrevskaya (2019). "Cephalonega, a new generic name, and the system of Vendian Proarticulata". Paleontological Journal. 53 (5): 447–454. doi:10.1134/S0031030119050046. S2CID 203853224.
^ Retallack, Gregory J. (January 2013). "Ediacaran life on land". Nature. 493 (7430): 89–92. doi:10.1038/nature11777. ISSN 0028-0836.
^ Xiao, Shuhai; Knauth, L. Paul (January 2013). "Fossils come in to land". Nature. 493 (7430): 28–29. doi:10.1038/nature11765. ISSN 0028-0836.
^ Xiao, Shuhai; Droser, Mary; Gehling, James G.; Hughes, Ian V.; Wan, Bin; Chen, Zhe; Yuan, Xunlai (March 2014). "Affirming life aquatic for the Ediacara biota in China and Australia: REPLY". Geology. 42 (3): e326. doi:10.1130/G35364Y.1. ISSN 1943-2682.
^ Seilacher, A. (January 2007). "The nature of vendobionts". Geological Society, London, Special Publications. 286 (1): 387–397. doi:10.1144/SP286.28. ISSN 0305-8719.
^ Hoyal Cuthill, Jennifer F.; Han, Jian (November 2018). Álvaro, Javier (ed.). "Cambrian petalonamid Stromatoveris phylogenetically links Ediacaran biota to later animals". Palaeontology. 61 (6): 813–823. doi:10.1111/pala.12393. ISSN 0031-0239.
^ Sperling, Erik; et al. (2008). "A placozoan affinity for Dickinsonia and the evolution of late Precambrian metazoan feeding modes". Geological Society of America. Abstracts with Programs. 40 (6): 508. Archived from the original on 28 February 2018. Retrieved 27 October 2008.
^ J.W. Valentine Dickinsonia as a polypoid organism Paleobiology, 18 (1992), pp. 378-382

[40] Unlike other species, D. costata has comparatively rounded body and fewer, wider segments / isomers.

[43] Dickinsonia menneri originally was identified as Vendomia but re-classified as Dickinsonia by Ivantsov (2007)^[2]

[44] D. menneri is a small organism up to 8 mm in length, and strongly resembles juvenile specimens of D. costata with its small number of isomers and well-marked head. D. menneri differs from juvenile D. costata by its slightly more elongated form.

[49] Dickinsonia tenuis strongly resembles D. costata, but differs from it by more narrow and numerous segments, sparingly lengthened oval form of the body.

[Hofmann1988-1] Hofmann, Hans J. (1988). "An alternative interpretation of the Ediacaran (Precambrian) chondrophore Chondroplon (Wade)". Alcheringa. 12 (4): 315–318. doi:10.1080/03115518808619130.

[Ivantsov2007-2] Ivantsov, A.Y. (2007a). "Small Vendian transversely articulated fossils". Paleontological Journal. 41 (2): 113–122. doi:10.1134/S0031030107020013. S2CID 86636748.

[:0-3] Gold, D.A.; Runnegar, B.; Gehling, J.G.; Jacobs, D.K. (2015). "Ancestral state reconstruction of ontogeny supports a bilaterian affinity for Dickinsonia". Evolution & Development. 17 (6): 315–397. doi:10.1111/ede.12168. PMID 26492825. S2CID 26099557.

[Pflug-4] Pflug (1973). "Zur fauna der Nama-Schichten in Südwest-Afrika. IV. Mikroscopische anatomie der petalo-organisme" [On the fauna of the Nama layers in southwest Africa. [part] IV. Microscopic anatomy of petaloorganisms]. Palaeontographica (B144): 166–202.

[Seilacher1992-5] Seilacher, Adolf (1992). "Vendobionta and Psammocorallia: Lost constructions of Precambrian evolution". Journal of the Geological Society, London. 149 (4): 607–613. Bibcode:1992JGSoc.149..607S. doi:10.1144/gsjgs.149.4.0607. S2CID 128681462. Retrieved 21 June 2007.

[McMenamin1998-6] McMenamin, M. (1998). The Garden of Ediacara. New York: Columbia University Press. ISBN 978-0-231-10559-0. OCLC 228271905.

[7] Dinosaurs: A visual encyclopedia. New York, NY: DK Publishing. 3 April 2018. ISBN 9781465469489.

[Bobrovskiy-8] Bobrovskiy, Ilya; Hope, Janet M.; Ivantsov, Andrey; Nettersheim, Benjamin J.; Hallmann, Christian; Brocks, Jochen J. (20 September 2018). "Ancient steroids establish the Ediacaran fossil Dickinsonia as one of the earliest animals". Science. 361 (6408): 1246–1249. Bibcode:2018Sci...361.1246B. doi:10.1126/science.aat7228. hdl:1885/230014. PMID 30237355.

[Love2021-9] Love, G.D.; Zumberge, J.A. (2021). "Emerging patterns in Proterozoic lipid biomarker records". Cambridge Elements. 361 (6408). doi:10.1017/9781108847117. ISBN 9781108847117.

[Rise2-10] Fedonkin, M.A.; Gehling, J.G.; Grey, K.; Narbonne, G.M.; Vickers-Rich, P. (2007). The Rise of Animals: Evolution and diversification of the kingdom Animalia. Johns Hopkins University Press. p. 326. ISBN 978-0-8018-8679-9.

[Ivantsov2012-11] Ivantsov, A. Yu (2012). "Becoming metamery and bilateral symmetry in Metazoa: Way of Proarticulata". Morphogenesis in the Individual and Historical Development: Symmetry and asymmetry: 16–17.

[:1-12] ^ ^a ^b ^c ^d ^e ^f Evans, Scott D.; Droser, Mary L.; Gehling, James G. (17 May 2017). Hejnol, Andreas (ed.). "Highly regulated growth and development of the Ediacara macrofossil Dickinsonia costata". PLoS One. 12 (5): e0176874. doi:10.1371/journal.pone.0176874. ISSN 1932-6203. PMC 5435172. PMID 28520741.

[Ivantsov2011-13] Ivantsov, A.Y. (2011). "Feeding traces of Proarticulata — the Vendian metazoa". Paleontological Journal. 45 (3): 237–248. doi:10.1134/S0031030111030063. S2CID 128741869.

[:3-14] ^ ^a ^b ^c ^d ^e ^f ^g ^h Ivantsov, Andrey Yu; Zakrevskaya, Maria (23 February 2023). "Body plan of Dickinsonia , the oldest mobile animals". Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 114 (1–2): 95–108. doi:10.1017/S175569102300004X. ISSN 1755-6910.

[:2-15] Dunn, Frances S.; Liu, Alexander G.; Donoghue, Philip C.J. (May 2018). "Ediacaran developmental biology". Biological Reviews. 93 (2): 914–932. doi:10.1111/brv.12379. ISSN 1464-7931. PMC 5947158. PMID 29105292.

[Ivantsov2004-16] Ivantsov, A.Y. (2004). "New Proarticulata from the Vendian of the Arkhangel'sk region" (PDF). Paleontological Journal. 38 (3): 247–253. Archived from the original (PDF) on 27 September 2007. Retrieved 27 September 2007.

[:5-17] Ivantsov, Andrey; Zakrevskaya, Maria; Nagovitsyn, Aleksey; Krasnova, Anna; Bobrovskiy, Ilya; Luzhnaya (Serezhnikova), Ekaterina (November 2020). "Intravital damage to the body of Dickinsonia (Metazoa of the late Ediacaran)". Journal of Paleontology. 94 (6): 1019–1033. doi:10.1017/jpa.2020.65. ISSN 0022-3360.

[:4-18] Evans, Scott D.; Hunt, Gene; Gehling, James G.; Sperling, Erik A.; Droser, Mary L. (January 2023). Rahman, Imran (ed.). "Species of Dickinsonia Sprigg from the Ediacaran of South Australia". Palaeontology. 66 (1). doi:10.1111/pala.12635. ISSN 0031-0239.

[Ivantsov2013-19] Ivantsov, A.Y. (2013). "Trace Fossils of Precambrian Metazoans "Vendobionta" and "Mollusks"". Stratigraphy and Geological Correlation. 21 (3): 252–264. Bibcode:2013SGC....21..252I. doi:10.1134/S0869593813030039. S2CID 128638405.

[Ivantsov_Malakhovskaya2002-20] Ivantsov, A.Y.; Malakhovskaya, Y.E. (2002). "Giant Traces of Vendian Animals" (PDF). Doklady Earth Sciences. 385 (6): 618–622. Archived from the original (PDF) on 4 July 2007. Retrieved 24 February 2008.

[21] Ivantsov, Andrey; Zakrevskaya, Maria (July 2022). "Dickinsonia : mobile and adhered". Geological Magazine. 159 (7): 1118–1133. doi:10.1017/S0016756821000194. ISSN 0016-7568.

[Sprigg1947-22] Sprigg, Reg C. (1947). "Early Cambrian (?) jellyfishes from the Flinders Ranges, South Australia" (PDF). Trans. R. Soc. S. Aust. 71: 212–24. Archived from the original (PDF) on 29 September 2007.

[23] Eig, Karsten. "EDIACARANS: THE FOSSILS THAT SHOULD NOT BE THERE". Adventures in geology - Karsten Eig.

[Fedonkin1983Ukr-24] Fedonkin, M. A. (1983). "Non-skeletal fauna of Podoloia, Dniester River valley". In Velikanov, V. A.; Assejeva, E. A.; Fedonkin, M. A. (eds.). The Vendian of the Ukraine (in Russian). Kiev: Naukova Dumka. pp. 128–139.

[Grazhdankin2004-25] Grazhdankin, Dima (2004). "Patterns of distribution in the Ediacaran biotas: facies versus biogeography and evolution" (PDF). Paleobiology. 30 (2): 203–221. doi:10.1666/0094-8373(2004)030<0203:PODITE>2.0.CO;2. S2CID 129376371.

[Maslov_2018-26] Maslov, A.V.; Podkovyrov, V.N.; Grazhdankin, D.V.; Kolesnikov, A.V. (2018). "Upper Vendian in the east, northeast and north of East European Platform: Depositional processes and biotic evolution". Litosfera. 18 (4): 520–542. doi:10.24930/1681-9004-2018-18-4-520-542.

[Kolesnikov_2018-27] Kolesnikov, A.V.; Liu, A.G.; Danelian, T.; Grazhdankin, D.V. (2018). "A reassessment of the problematic Ediacaran genus Orbisiana Sokolov 1976". Precambrian Research. 316: 197–205. Bibcode:2018PreR..316..197K. doi:10.1016/j.precamres.2018.08.011. S2CID 134213721.

[28] Wang, Xiao-Peng; Chen, Zhe; Pang, Ke; Zhou, Chuan-Ming; Xiao, Shuhai; Wan, Bin; Yuan, Xun-Lai (2021). "Dickinsonia from the Ediacaran Dengying Formation in the Yangtze Gorges area, South China". Palaeoworld. 30 (4): 602–609. doi:10.1016/j.palwor.2021.01.002. S2CID 234254399.

[Ivantsov2009-29] Ivantsov, A.Y. (2009). "A new reconstruction of Kimberella, a problematic Vendian Metazoan". Paleontological Journal. 43 (6): 601–611. doi:10.1134/S003103010906001X. S2CID 85676210.

[Gehling1999-30] Gehling, J.G. (1999). "Microbial mats in terminal Proterozoic siliciclastics; Ediacaran death masks". PALAIOS. 14 (1): 40–57. Bibcode:1999Palai..14...40G. doi:10.2307/3515360. JSTOR 3515360.

[Serezhnikova2011-31] Serezhnikova Z, E.A. (2011). "Microbial Binding as a Probable Cause of Taphonomic Variability of Vendian Fossils: Carbonate Casting?". Advances in Stromatolite Geobiology. Lecture Notes in Earth Sciences. Vol. 131. pp. 525–535. doi:10.1007/978-3-642-10415-2_31. ISBN 978-3-642-10414-5.

[Runnegar1982-32] Runnegar, Bruce (1982). "Oxygen requirements, biology and phylogenetic significance of the late Precambrian worm Dickinsonia, and the evolution of the burrowing habit". Alcheringa: An Australasian Journal of Palaeontology. 6 (3): 223–239. doi:10.1080/03115518208565415.

[Grazhdankin20042-33] Grazhdankin, Dima (2004). "Patterns of distribution in the Ediacaran biotas: facies versus biogeography and evolution" (PDF). Paleobiology. 30 (2): 203–221. doi:10.1666/0094-8373(2004)030<0203:PODITE>2.0.CO;2. S2CID 129376371.

[Keller_Fedonkin_1976-34] Keller, B.M.; Fedonkin, M.A. (1976). "New records of fossils in the Valdaian group of the precambrian on the Syuz'ma River" (PDF). Izvestiya Akademii Nauk SSR. Seriya Geologicheskaya (in Russian). 3: 38–44. Archived from the original (PDF) on 27 September 2007.

[Keller_Fedonkin_1977-35] Keller, B. M.; Fedonkin, M. A. (1977). "New organic fossil finds in the Precambrian Valday series along the Syuz'ma River". International Geology Review. 19 (8): 924–930. Bibcode:1977IGRv...19..924K. doi:10.1080/00206817709471091.

[Gehling_Droser2013-36] Gehling, J.G.; Droser, M.L. (2013). "How well do fossil assemblages of the Ediacara Biota tell time?". Geology. 41 (4): 447–450. Bibcode:2013Geo....41..447G. doi:10.1130/G33881.1.

[Wade1971-37] Wade, M. (1971). "Bilateral Precambrian Chondrophores from the Ediacara Fauna, South Australia". Proceedings of the Royal Society of Victoria. 84 (1): 183–188.

[Evans2023-38] Evans, S.D.; Hunt, G.; Gehling, J.G.; Sperling, E. A.; Droser, M.L. (2023). "Species of Dickinsonia sprigg from the Ediacaran of South Australia". Palaeontology. 66 (1–21): e12635. doi:10.1111/pala.12635.

[Wade1972-39] Wade, M. (1972). "Dickinsonia: Polychaete worms from the late Precambrian Ediacara fauna, South Australia". Mem. Queensl. Mus. 16 (2): 171–190.

[Sprigg19472-41] Sprigg, Reg C. (1947a). "Early Cambrian (?) jellyfishes from the Flinders Ranges, South Australia" (PDF). Trans. R. Soc. S. Aust. 71: 212–24. Archived from the original (PDF) on 29 September 2007.

[Glaessner1966-42] Glaessner, M.F.; Wade, M. (1966). "The late Precambrian fossils from Ediacara, South Australia" (PDF). Palaeontology. 9 (4): 599.

[Keller_Fedonkin_19762-45] Keller, B.M.; Fedonkin, M.A. (1976). "New records of fossils in the Valdaian group of the precambrian on the Syuz'ma River" (PDF). Izvestiya Akademii Nauk SSR. Seriya Geologicheskaya (in Russian). 3: 38–44. Archived from the original (PDF) on 27 September 2007.

[Sprigg1949-46] Sprigg, R.C. (1949). "Early Cambrian "jellyfishes" of Ediacara, South Australia, and Mount John, Kimberley District, Western Australia" (PDF). Transactions of the Royal Society of South Australia. 73: 72–99.{{cite journal}}: CS1 maint: url-status (link)^{[permanent dead link]}

[Jenkins1992-47] Jenkins, R.J.F. (1992). "Functional and ecological aspects of Ediacarian assemblages". In Lipps, J.; Signor, P.W. (eds.). Origin and Early Evolution of the Metazoa. New York, NY: Springer. pp. 131–176. ISBN 978-0-306-44067-0. OCLC 231467647.

[Harrington_Moore1955-48] Harrington, N.J.; Moore, R.C. (1955). "Kansas Pennsylvanian and other jellyfishes". Bull. Kansas Geol. Surv. 114 (5): 153–163.

[50] Pandey, S.K.; Ahmad, Shamim; Sharma, Mukund (9 March 2023). "Dickinsonia tenuis reported by Retallack et al. 2021 is not a fossil, instead an impression of an extant 'fallen beehive'". Journal of the Geological Society of India. 99 (3): 311–316. doi:10.1007/s12594-023-2312-2. ISSN 0974-6889.

[Ivantsov_2019-51] Ivantsov, A.Y.; Fedonkin, M.A.; Nagovitsyn, A.L.; M.A. Zakrevskaya (2019). "Cephalonega, a new generic name, and the system of Vendian Proarticulata". Paleontological Journal. 53 (5): 447–454. doi:10.1134/S0031030119050046. S2CID 203853224.

[52] Retallack, Gregory J. (January 2013). "Ediacaran life on land". Nature. 493 (7430): 89–92. doi:10.1038/nature11777. ISSN 0028-0836.

[53] Xiao, Shuhai; Knauth, L. Paul (January 2013). "Fossils come in to land". Nature. 493 (7430): 28–29. doi:10.1038/nature11765. ISSN 0028-0836.

[54] Xiao, Shuhai; Droser, Mary; Gehling, James G.; Hughes, Ian V.; Wan, Bin; Chen, Zhe; Yuan, Xunlai (March 2014). "Affirming life aquatic for the Ediacara biota in China and Australia: REPLY". Geology. 42 (3): e326. doi:10.1130/G35364Y.1. ISSN 1943-2682.

[55] Seilacher, A. (January 2007). "The nature of vendobionts". Geological Society, London, Special Publications. 286 (1): 387–397. doi:10.1144/SP286.28. ISSN 0305-8719.

[56] Hoyal Cuthill, Jennifer F.; Han, Jian (November 2018). Álvaro, Javier (ed.). "Cambrian petalonamid Stromatoveris phylogenetically links Ediacaran biota to later animals". Palaeontology. 61 (6): 813–823. doi:10.1111/pala.12393. ISSN 0031-0239.

[Sperling2008-57] Sperling, Erik; et al. (2008). "A placozoan affinity for Dickinsonia and the evolution of late Precambrian metazoan feeding modes". Geological Society of America. Abstracts with Programs. 40 (6): 508. Archived from the original on 28 February 2018. Retrieved 27 October 2008.

[58] J.W. Valentine Dickinsonia as a polypoid organism Paleobiology, 18 (1992), pp. 378-382

[1]

[2]

[3]

[4]

[5]

[6]

[7]

[8]

[9]

[10]

[11]

[12]

[13]

[14]

[15]

[16]

[17]

[18]

[19]

[20]

[21]

[22]

[23]

[24]

[25]

[26]

[27]

[28]

[29]

[30]

[31]

[32]

[33]

[34]

[35]

[36]

[37]

[38]

[39]

[A]

[40]

[41]

[B]

[C]

[42]

[43]

[44]

[45]

[D]

[46]

[47]

[48]

[49]

[50]

[51]

[52]

[53]

[54]

@@ Line 1: / Line 1: @@
 {{Short description|Extinct genus of early animals}}
-{{distinguish|Dicksonia}}
+{{distinguish|Dicksonia}}{{use dmy dates|date=April 2024}}
 {{Automatic taxobox
 | fossil_range = Late [[Ediacaran]], {{fossil range|567|550}}
@@ Line 12: / Line 12: @@
 | subdivision_ranks= Species
 | subdivision =
-* ''D. costata'' {{small|Sprigg, 1947}}
+* ''D.&nbsp;costata'' {{small|(Sprigg, 1947)}}
-* ''D. menneri'' {{small|Keller, 1976}}
+* ''D.&nbsp;menneri'' {{small|(Keller, 1976)}}
-* ''D. tenuis'' {{small|Glaessner & Wade, 1966}}
+* ''D.&nbsp;tenuis'' {{small|(Glaessner & Wade, 1966)}}
 | synonyms =
 {{collapsible list|bullets = true
- |title = <small>Genus Synonymy</small>
+ |title = <small>Genus synonymy</small>
-| ''Chondroplon''? {{small|Wade, 1971}}<ref name=Hofmann1988>{{cite journal | author=Hofmann, Hans J. | year=1988 | title=An alternative interpretation of the Ediacaran (Precambrian) chondrophore ''Chondroplon'' Wade | journal=Alcheringa | pages=315–318 | volume=12
+| ''Chondroplon''? {{small|(Wade, 1971)}}<ref name=Hofmann1988>{{cite journal | last=Hofmann | first=Hans J. | year=1988 | title=An alternative interpretation of the Ediacaran (Precambrian) chondrophore ''Chondroplon'' {{small|(Wade)}} | journal=Alcheringa | volume=12 | issue=4 | pages=315–318 | doi=10.1080/03115518808619130}}</ref>
- | issue=4 | doi=10.1080/03115518808619130}}</ref>
 | ''Papilionata'' {{small|Sprigg, 1947}}
-| ''Vendomia'' {{small|Keller, 1976}}<ref name="Ivantsov20072"/>
+| ''Vendomia'' {{small|Keller, 1976}}<ref name=Ivantsov2007/>
 }}
 {{collapsible list|bullets = true
- |title = <small>''D. costata'' Synonymy</small>
+ |title = <small>''D.&nbsp;costata'' synonymy</small>
- |''Papilionata eyrei'' {{small|Sprigg, 1947}}
+ |''Papilionata eyrei'' {{small|(Sprigg, 1947)}}
- |''D. minima'' {{small|Sprigg, 1949}}
+ |''D.&nbsp;minima'' {{small|(Sprigg, 1949}}
- |''D. elongata'' {{small|Glaessner & Wade, 1966}}
+ |''D.&nbsp;elongata'' {{small|(Glaessner & Wade, 1966)}}
- |''D. spriggi'' {{small|Harrington & Moore, 1955}}
+ |''D.&nbsp;spriggi'' {{small|(Harrington & Moore, 1955)}}
 }}
 {{collapsible list|bullets = true
- |title = <small>''D. tenuis'' Synonymy</small>
+ |title = <small>''D.&nbsp;tenuis'' synonymy</small>
- |''D. brachina'' {{small|Wade, 1972}}
+ |''D.&nbsp;brachina'' {{small|(Wade, 1972)}}
- |''D. lissa'' {{small|Wade, 1972}}
+ |''D.&nbsp;lissa'' {{small|(Wade, 1972)}}
- |''D. rex'' {{small|Jenkins, 1992}}
+ |''D.&nbsp;rex'' {{small|(Jenkins, 1992)}}
 }}
 }}
-'''''Dickinsonia''''' is a genus of extinct organism, most likely an [[animal]], that lived during the late [[Ediacaran]] period in what is now Australia, China, Russia and Ukraine. It is one of the best known members of the [[Ediacaran biota]]. The individual ''Dickinsonia'' typically resembles a bilaterally symmetrical ribbed oval. Its [[Affinity (taxonomy)|affinities]] are presently unknown; its mode of growth has been considered consistent with a stem-group [[bilateria]]n affinity,<ref name=":0">{{Cite journal |last1=Gold |first1=D. A. |last2=Runnegar |first2=B. |last3=Gehling |first3=J. G. |last4=Jacobs |first4=D. K. |year=2015 |title=Ancestral state reconstruction of ontogeny supports a bilaterian affinity for Dickinsonia |journal=Evolution & Development |volume=17 |issue=6 |pages=315–397 |doi=10.1111/ede.12168 |pmid=26492825 |s2cid=26099557}}</ref> though various other affinities have been proposed.<ref name="Pflug">{{Cite journal |author=Pflug |year=1973 |title=Zur fauna der Nama-Schichten in Südwest-Afrika. IV. Mikroscopische anatomie der petalo-organisme |journal=Palaeontographica |issue=B144 |pages=166–202}}</ref><ref name="Seilacher1992">{{Cite journal |author=Seilacher, Adolf |author-link=Adolf Seilacher |year=1992 |title=Vendobionta and Psammocorallia: lost constructions of Precambrian evolution |url=http://jgs.lyellcollection.org/cgi/content/abstract/149/4/607 |journal=Journal of the Geological Society, London |volume=149 |issue=4 |pages=607–613 |bibcode=1992JGSoc.149..607S |doi=10.1144/gsjgs.149.4.0607 |s2cid=128681462 |access-date=2007-06-21}}</ref><ref name="McMenamin1998">{{Cite book |author=McMenamin M. |title=The Garden of Ediacara |publisher=Columbia University Press |year=1998 |isbn=978-0-231-10559-0 |location=New York |oclc=228271905}} It lived during the late Ediacaran (part of Precambrian). {{Cite book |title=Dinosaurs a visual encyclopedia |date=2018-04-03 |publisher=DK Publishing, Inc |isbn=9781465469489 |location=New York}}</ref> The discovery of [[cholesterol]] molecules in fossils of ''Dickinsonia'' lends support to the idea that ''Dickinsonia'' was an animal,<ref name="Bobrovskiy">{{cite journal |last1=Bobrovskiy |first1=Ilya |last2=Hope |first2=Janet M. |last3=Ivantsov |first3=Andrey |last4=Nettersheim |first4=Benjamin J. |last5=Hallmann |first5=Christian |last6=Brocks |first6=Jochen J. |date=20 September 2018 |title=Ancient steroids establish the Ediacaran fossil Dickinsonia as one of the earliest animals |journal=Science |volume=361 |issue=6408 |pages=1246–1249 |bibcode=2018Sci...361.1246B |doi=10.1126/science.aat7228 |pmid=30237355 |doi-access=free|hdl=1885/230014 |hdl-access=free }}</ref> though these results have been questioned.<ref name="Love2021">{{cite journal |last1=Love |first1=G.D. |last2=Zumberge |first2=J.A. |date=2021 |title=Emerging patterns in Proterozoic lipid biomarker records |journal=Cambridge Elements |volume=361 |issue=6408 |doi=10.1017/9781108847117|isbn=9781108847117 }}</ref>
+'''''Dickinsonia''''' is a genus of extinct organism, most likely an [[animal]], that lived during the late [[Ediacaran]] period in what is now Australia, China, Russia, and Ukraine. It is one of the best known members of the [[Ediacaran biota]]. The individual ''Dickinsonia'' typically resembles a bilaterally symmetrical ribbed oval. Its [[Affinity (taxonomy)|affinities]] are presently unknown; its mode of growth has been considered consistent with a stem-group [[bilateria]]n affinity,<ref name=":0">{{cite journal |last1=Gold |first1=D.A. |last2=Runnegar |first2=B. |last3=Gehling |first3=J.G. |last4=Jacobs |first4=D.K. |year=2015 |title=Ancestral state reconstruction of ontogeny supports a bilaterian affinity for Dickinsonia |journal=Evolution & Development |volume=17 |issue=6 |pages=315–397 |doi=10.1111/ede.12168 |pmid=26492825 |s2cid=26099557}}</ref> though various other affinities have been proposed.<ref name=Pflug>{{cite journal |last=Pflug |first= |year=1973 |title=Zur fauna der Nama-Schichten in Südwest-Afrika. IV.&nbsp;Mikroscopische anatomie der petalo-organisme |trans-title=On the fauna of the Nama layers in southwest Africa. {{nobr|{{grey|[part]}} IV. Microscopic}} anatomy of petaloorganisms |journal=Palaeontographica |issue=B144 |pages=166–202}}</ref><ref name="Seilacher1992">{{cite journal |last=Seilacher |first=Adolf |author-link=Adolf Seilacher |year=1992 |title=Vendobionta and Psammocorallia: Lost constructions of Precambrian evolution |journal=Journal of the Geological Society, London |volume=149 |issue=4 |pages=607–613 |bibcode=1992JGSoc.149..607S |doi=10.1144/gsjgs.149.4.0607 |s2cid=128681462 |url=http://jgs.lyellcollection.org/cgi/content/abstract/149/4/607 |access-date=2007-06-21}}</ref><ref name=McMenamin1998>{{cite book |last=McMenamin |first=M. |year=1998 |title=The Garden of Ediacara |publisher=Columbia University Press |isbn=978-0-231-10559-0 |location=New York |oclc=228271905}}</ref> It lived during the late Ediacaran (final part of Precambrian).<ref>{{cite book |title=Dinosaurs: A visual encyclopedia |date=2018-04-03 |df=dmy-all |place=New York, NY |publisher=DK Publishing |isbn=9781465469489 }}</ref> The discovery of [[cholesterol]] molecules in fossils of ''Dickinsonia'' lends support to the idea that ''Dickinsonia'' was an animal,<ref name=Bobrovskiy>{{cite journal |last1=Bobrovskiy |first1=Ilya |last2=Hope |first2=Janet M. |last3=Ivantsov |first3=Andrey |last4=Nettersheim |first4=Benjamin J. |last5=Hallmann |first5=Christian |last6=Brocks |first6=Jochen J. |date=20 September 2018 |title=Ancient steroids establish the Ediacaran fossil Dickinsonia as one of the earliest animals |journal=Science |volume=361 |issue=6408 |pages=1246–1249 |bibcode=2018Sci...361.1246B |doi=10.1126/science.aat7228 |doi-access=free |pmid=30237355 |hdl=1885/230014 |hdl-access=free }}</ref> though these results have been questioned.<ref name=Love2021>{{cite journal |last1=Love |first1=G.D. |last2=Zumberge |first2=J.A. |year=2021 |title=Emerging patterns in Proterozoic lipid biomarker records |journal=Cambridge Elements |volume=361 |issue=6408 |doi=10.1017/9781108847117 |isbn=9781108847117 }}</ref>
 == Description ==
-''Dickinsonia'' fossils are known only in the form of imprints and casts in sandstone beds. The specimens found range from a few [[millimetre]]s to about {{convert|1.4|m}} in length, and from a fraction of a millimetre to a few millimetres thick.<ref name="Rise2">{{cite book |last1=Fedonkin |first1=2 M. A. |url=https://books.google.com/books?id=OFKG6SmPNuUC |title=The Rise of Animals. Evolution and Diversification of the Kingdom Animalia |last2=Gehling |first2=J. G. |last3=Grey |first3=K. |last4=Narbonne |first4=G. M. |last5=Vickers-Rich |first5=P. |publisher=Johns Hopkins University Press |year=2007 |isbn=978-0-8018-8679-9 |pages=326}}</ref> They are nearly bilaterally symmetric, segmented, round or oval in outline, slightly expanded to one end (i.e. egg-shaped outline). The rib-like segments are radially inclined towards the wide and narrow ends, and the width and length of the segments increases towards the wide end of the fossil.<ref name="Ivantsov20072">{{cite journal |last=Ivantsov |first=A. Y. |year=2007 |title=Small Vendian transversely articulated fossils |url=https://www.academia.edu/2352394 |journal=Paleontological Journal |volume=41 |issue=2 |pages=113–122 |doi=10.1134/S0031030107020013 |s2cid=86636748}}</ref><ref name="Ivantsov2012p">{{cite journal |author=Ivantsov, A. Yu |year=2012 |title=Becoming metamery and bilateral symmetry in Metazoa: way of Proarticulata |journal=Morphogenesis in the Individual and Historical Development: Symmetry and Asymmetry. |pages=16–17}}</ref> The body is divided into two by a midline ridge or groove,<ref name="Ivantsov20072"/><ref name="Ivantsov2012p" /><ref name=":1">{{Cite journal |last1=Evans |first1=Scott D. |last2=Droser |first2=Mary L. |last3=Gehling |first3=James G. |date=2017-05-17 |editor-last=Hejnol |editor-first=Andreas |title=Highly regulated growth and development of the Ediacara macrofossil Dickinsonia costata |journal=PLOS ONE |language=en |volume=12 |issue=5 |pages=e0176874 |doi=10.1371/journal.pone.0176874 |issn=1932-6203 |pmc=5435172 |pmid=28520741 |doi-access=free }}</ref> except for a single unpaired segment at one end, dubbed the "anterior most unit" suggested to represent the front of the organism.<ref name=":1" /> Is it disputed whether the segments are offset from each other following [[glide reflection]], and are thus isomers,<ref name="Ivantsov20072"/><ref name="Ivantsov2012p" /><ref name="Ivantsov20113">{{Cite journal |last1=Ivantsov |first1=A. Y. |year=2011 |title=Feeding traces of Proarticulata — the Vendian metazoa |url=https://www.academia.edu/2394670 |journal=Paleontological Journal |volume=45 |issue=3 |pages=237–248 |doi=10.1134/S0031030111030063 |s2cid=128741869}}</ref><ref name=":3" /> or whether the segments are symmetric across the midline, and thus follow true [[bilateral symmetry]], as the specimens displaying the offset may be the result of [[Taphonomy|taphonomic distortion]].<ref name=":1" /><ref name=":2">{{Cite journal |last1=Dunn |first1=Frances S. |last2=Liu |first2=Alexander G. |last3=Donoghue |first3=Philip C. J. |date=May 2018 |title=Ediacaran developmental biology |journal=Biological Reviews |language=en |volume=93 |issue=2 |pages=914–932 |doi=10.1111/brv.12379 |issn=1464-7931 |pmc=5947158 |pmid=29105292}}</ref>  The number of segments/isomer pairs varies from 12 in smaller individuals to 74 in the largest Australian specimens.<ref name=":2" />
+''Dickinsonia'' fossils are known only in the form of imprints and casts in sandstone beds. The specimens found range from a few [[millimetre]]s to about {{convert|1.4|m}} in length, and from a fraction of a millimetre to a few millimetres thick.<ref name=Rise2>{{cite book |last1=Fedonkin |first1=M.A. |last2=Gehling |first2=J.G. |last3=Grey |first3=K. |last4=Narbonne |first4=G.M. |last5=Vickers-Rich |first5=P. |year=2007 |title=The Rise of Animals: Evolution and diversification of the kingdom Animalia |publisher=Johns Hopkins University Press |isbn=978-0-8018-8679-9 |page=326 |url=https://books.google.com/books?id=OFKG6SmPNuUC }}</ref> They are nearly bilaterally symmetric, segmented, round or oval in outline, slightly expanded to one end (i.e. egg-shaped outline). The rib-like segments are radially inclined towards the wide and narrow ends, and the width and length of the segments increases towards the wide end of the fossil.<ref name=Ivantsov2007>{{cite journal |last=Ivantsov |first=A.Y. |year=2007a |title=Small Vendian transversely articulated fossils |journal=Paleontological Journal |volume=41 |issue=2 |pages=113–122 |doi=10.1134/S0031030107020013 |s2cid=86636748 |url=https://www.academia.edu/2352394 }}</ref><ref name=Ivantsov2012>{{cite journal |last=Ivantsov |first=A. Yu |year=2012 |title=Becoming metamery and bilateral symmetry in Metazoa: Way of Proarticulata |journal=Morphogenesis in the Individual and Historical Development: Symmetry and asymmetry |pages=16–17}}</ref> The body is divided into two by a midline ridge or groove,<ref name=Ivantsov2007/><ref name=Ivantsov2012/><ref name=":1">{{cite journal |last1=Evans |first1=Scott D. |last2=Droser |first2=Mary L. |last3=Gehling |first3=James G. |date=2017-05-17 |editor-last=Hejnol |editor-first=Andreas |title=Highly regulated growth and development of the Ediacara macrofossil Dickinsonia costata |journal=PLoS One |lang=en |volume=12 |issue=5 |page=e0176874 |doi=10.1371/journal.pone.0176874 |doi-access=free |issn=1932-6203 |pmc=5435172 |pmid=28520741 }}</ref> except for a single unpaired segment at one end, dubbed the "anterior most unit" suggested to represent the front of the organism.<ref name=":1" /> Is it disputed whether the segments are offset from each other following [[glide reflection]], and are thus isomers,<ref name=Ivantsov2007/><ref name=Ivantsov2012/><ref name=Ivantsov2011>{{cite journal |last1=Ivantsov |first1=A.Y. |year=2011 |title=Feeding traces of Proarticulata — the Vendian metazoa |journal=Paleontological Journal |volume=45 |issue=3 |pages=237–248 |doi=10.1134/S0031030111030063 |s2cid=128741869 |url=https://www.academia.edu/2394670}}</ref><ref name=":3"/> or whether the segments are symmetric across the midline, and thus follow true [[bilateral symmetry]], as the specimens displaying the offset may be the result of [[Taphonomy|taphonomic distortion]].<ref name=":1" /><ref name=":2">{{cite journal |last1=Dunn |first1=Frances S. |last2=Liu |first2=Alexander G. |last3=Donoghue |first3=Philip C.J. |date=May 2018 |title=Ediacaran developmental biology |journal=Biological Reviews |lang=en |volume=93 |issue=2 |pages=914–932 |doi=10.1111/brv.12379 |issn=1464-7931 |pmc=5947158 |pmid=29105292}}</ref>  The number of segments/isomer pairs varies from 12 in smaller individuals to 74 in the largest Australian specimens.<ref name=":2"/>
-The body of ''Dickinsonia'' is suggested to have been sack-like, with the outer layer being made of a resistant but unmineralised material.<ref name=":3">{{Cite journal |last1=Ivantsov |first1=Andrey Yu |last2=Zakrevskaya |first2=Maria |date=2023-02-23 |title=Body plan of Dickinsonia , the oldest mobile animals |url=https://www.cambridge.org/core/product/identifier/S175569102300004X/type/journal_article |journal=Earth and Environmental Science Transactions of the Royal Society of Edinburgh |volume=114 |issue=1–2 |language=en |pages=95–108 |doi=10.1017/S175569102300004X |issn=1755-6910}}</ref> Some specimens from Russia show the presence of branched internal structures.<ref name="Ivantsov2004">{{cite journal |author=Ivantsov, A. Y. |year=2004 |title=New Proarticulata from the Vendian of the Arkhangel'sk Region |url=http://vend.paleo.ru/pub/Ivantsov_2004_eng.pdf |url-status=dead |journal=Paleontological Journal |volume=38 |issue=3 |pages=247–253 |archive-url=https://web.archive.org/web/20070927022709/http://vend.paleo.ru/pub/Ivantsov_2004_eng.pdf |archive-date=2007-09-27 |access-date=2007-09-27}}</ref><ref name=":3" /> Some authors have suggested that the underside of the body bore [[Cilium|cilia]], as well as infolded pockets.<ref name=":3" />
+The body of ''Dickinsonia'' is suggested to have been sack-like, with the outer layer being made of a resistant but unmineralised material.<ref name=":3">{{cite journal |last1=Ivantsov |first1=Andrey Yu |last2=Zakrevskaya |first2=Maria |date=2023-02-23 |title=Body plan of Dickinsonia , the oldest mobile animals |url=https://www.cambridge.org/core/product/identifier/S175569102300004X/type/journal_article |journal=Earth and Environmental Science Transactions of the Royal Society of Edinburgh |volume=114 |issue=1–2 |language=en |pages=95–108 |doi=10.1017/S175569102300004X |issn=1755-6910}}</ref> Some specimens from Russia show the presence of branched internal structures.<ref name=Ivantsov2004>{{cite journal |last=Ivantsov |first=A.Y. |year=2004 |title=New Proarticulata from the Vendian of the Arkhangel'sk region |url-status=dead |journal=Paleontological Journal |volume=38 |issue=3 |pages=247–253 |url=http://vend.paleo.ru/pub/Ivantsov_2004_eng.pdf |access-date=2007-09-27 |archive-url=https://web.archive.org/web/20070927022709/http://vend.paleo.ru/pub/Ivantsov_2004_eng.pdf |archive-date=2007-09-27}}</ref><ref name=":3"/> Some authors have suggested that the underside of the body bore [[Cilium|cilia]], as well as infolded pockets.<ref name=":3"/>
-''Dickinsonia'' is suggested to have grown by adding a new pair of segments/isomers at the end opposite the unpaired "anterior most unit".<ref name=":1" /><ref name=":5">{{Cite journal |last1=Ivantsov |first1=Andrey |last2=Zakrevskaya |first2=Maria |last3=Nagovitsyn |first3=Aleksey |last4=Krasnova |first4=Anna |last5=Bobrovskiy |first5=Ilya |last6=Luzhnaya (Serezhnikova) |first6=Ekaterina |date=November 2020 |title=Intravital damage to the body of Dickinsonia (Metazoa of the late Ediacaran) |url=https://www.cambridge.org/core/product/identifier/S0022336020000657/type/journal_article |journal=Journal of Paleontology |language=en |volume=94 |issue=6 |pages=1019–1033 |doi=10.1017/jpa.2020.65 |issn=0022-3360}}</ref> ''Dickinsonia'' probably exhibited [[indeterminate growth]] (having no maximum size), though it is suggested that the addition of new segments slowed down later in growth.<ref name=":4">{{Cite journal |last1=Evans |first1=Scott D. |last2=Hunt |first2=Gene |last3=Gehling |first3=James G. |last4=Sperling |first4=Erik A. |last5=Droser |first5=Mary L. |date=January 2023 |editor-last=Rahman |editor-first=Imran |title=Species of Dickinsonia Sprigg from the Ediacaran of South Australia |journal=Palaeontology |language=en |volume=66 |issue=1 |doi=10.1111/pala.12635 |issn=0031-0239|doi-access=free }}</ref> Deformed specimens from Russia indicate that individuals of ''Dickinsonia'' could regenerate after being damaged.<ref name=":5" />
+''Dickinsonia'' is suggested to have grown by adding a new pair of segments/isomers at the end opposite the unpaired "anterior most unit".<ref name=":1"/><ref name=":5">{{cite journal |last1=Ivantsov |first1=Andrey |last2=Zakrevskaya |first2=Maria |last3=Nagovitsyn |first3=Aleksey |last4=Krasnova |first4=Anna |last5=Bobrovskiy |first5=Ilya |last6=Luzhnaya (Serezhnikova) |first6=Ekaterina |date=November 2020 |title=Intravital damage to the body of Dickinsonia (Metazoa of the late Ediacaran) |url=https://www.cambridge.org/core/product/identifier/S0022336020000657/type/journal_article |journal=Journal of Paleontology |language=en |volume=94 |issue=6 |pages=1019–1033 |doi=10.1017/jpa.2020.65 |issn=0022-3360}}</ref> ''Dickinsonia'' probably exhibited [[indeterminate growth]] (having no maximum size), though it is suggested that the addition of new segments slowed down later in growth.<ref name=":4">{{cite journal |last1=Evans |first1=Scott D. |last2=Hunt |first2=Gene |last3=Gehling |first3=James G. |last4=Sperling |first4=Erik A. |last5=Droser |first5=Mary L. |date=January 2023 |editor-last=Rahman |editor-first=Imran |title=Species of Dickinsonia Sprigg from the Ediacaran of South Australia |journal=Palaeontology |language=en |volume=66 |issue=1 |doi=10.1111/pala.12635 |issn=0031-0239|doi-access=free }}</ref> Deformed specimens from Russia indicate that individuals of ''Dickinsonia'' could regenerate after being damaged.<ref name=":5"/>
-{{gallery|Dickinsonia costata Ontogeny.jpg|Ontogeny of ''Dickinsonia costata'' following the glide reflection interpretation|Dickinsonia growth.png|Growth of ''D. costata'' under bilateral symmetry interpretation|Dickinsonia species 2.png|Diagram of various ''Dickinsonia'' species|Dickinsonia species.png|Diagram of various ''Dickinsonia'' species (cont)|Dickinsonia internal anatomy.svg|Diagram of branched internal structures observed in Russian specimens|width=225|height=250|lines=|align=center}}
+{{gallery|Dickinsonia costata Ontogeny.jpg|Ontogeny of ''Dickinsonia costata'' following the glide reflection interpretation|Dickinsonia growth.png|Growth of ''D.&nbsp;costata'' under bilateral symmetry interpretation|Dickinsonia species 2.png|Diagram of various ''Dickinsonia'' species|Dickinsonia species.png|Diagram of various ''Dickinsonia'' species (cont)|Dickinsonia internal anatomy.svg|Diagram of branched internal structures observed in Russian specimens|width=225|height=250|lines=|align=center}}
 == Ecology ==
-''Dickinsonia'' is suggested to have been a mobile marine organism that lived on the seafloor and fed by consuming [[microbial mat]]s growing on the seabed using structures present on its underside. ''Dickinsonia''-shaped [[trace fossil]]s, presumed to represent feeding impressions, sometimes found in chains demonstrating this behaviour have been observed.<ref name=":3" /> These trace fossils have been assigned to the genus ''[[Epibaion]]''.<ref name="Ivantsov20113"/><ref name="Ivantsov20132">{{cite journal |last=Ivantsov |first=A.Y. |year=2013 |title=Trace Fossils of Precambrian Metazoans "Vendobionta" and "Mollusks" |url=https://www.academia.edu/3757185 |journal=Stratigraphy and Geological Correlation |volume=21 |issue=3 |pages=252–264 |bibcode=2013SGC....21..252I |doi=10.1134/S0869593813030039 |s2cid=128638405}}</ref><ref name="Ivantsov_Malakhovskaya2002">{{cite journal |last1=Ivantsov |first1=A.Y. |last2=Malakhovskaya |first2=Y.E. |year=2002 |title=Giant Traces of Vendian Animals |url=http://vend.paleo.ru/pub/Ivantsov_et_Malakhovskaya_2002-e.pdf |url-status=dead |journal=Doklady Earth Sciences |volume=385 |issue=6 |pages=618–622 |archive-url=https://web.archive.org/web/20070704183947/http://vend.paleo.ru/pub/Ivantsov_et_Malakhovskaya_2002-e.pdf |archive-date=2007-07-04 |access-date=2008-02-24}}</ref> A 2022 study suggested that ''Dickinsonia'' temporarily adhered itself to the seafloor by the use of [[mucus]], which may have been an adaptation to living in very shallow water environments.<ref>{{Cite journal |last1=Ivantsov |first1=Andrey |last2=Zakrevskaya |first2=Maria |date=July 2022 |title=Dickinsonia : mobile and adhered |url=https://www.cambridge.org/core/product/identifier/S0016756821000194/type/journal_article |journal=Geological Magazine |language=en |volume=159 |issue=7 |pages=1118–1133 |doi=10.1017/S0016756821000194 |issn=0016-7568}}</ref>
+''Dickinsonia'' is suggested to have been a mobile marine organism that lived on the seafloor and fed by consuming [[microbial mat]]s growing on the seabed using structures present on its underside. ''Dickinsonia''-shaped [[trace fossil]]s, presumed to represent feeding impressions, sometimes found in chains demonstrating this behaviour have been observed.<ref name=":3"/> These trace fossils have been assigned to the genus ''[[Epibaion]]''.<ref name=Ivantsov2011/><ref name=Ivantsov2013>{{cite journal |last=Ivantsov |first=A.Y. |year=2013 |title=Trace Fossils of Precambrian Metazoans "Vendobionta" and "Mollusks" |url=https://www.academia.edu/3757185 |journal=Stratigraphy and Geological Correlation |volume=21 |issue=3 |pages=252–264 |bibcode=2013SGC....21..252I |doi=10.1134/S0869593813030039 |s2cid=128638405}}</ref><ref name=Ivantsov_Malakhovskaya2002>{{cite journal |last1=Ivantsov |first1=A.Y. |last2=Malakhovskaya |first2=Y.E. |year=2002 |title=Giant Traces of Vendian Animals |url=http://vend.paleo.ru/pub/Ivantsov_et_Malakhovskaya_2002-e.pdf |url-status=dead |journal=Doklady Earth Sciences |volume=385 |issue=6 |pages=618–622 |archive-url=https://web.archive.org/web/20070704183947/http://vend.paleo.ru/pub/Ivantsov_et_Malakhovskaya_2002-e.pdf |archive-date=2007-07-04 |access-date=2008-02-24}}</ref> A 2022 study suggested that ''Dickinsonia'' temporarily adhered itself to the seafloor by the use of [[mucus]], which may have been an adaptation to living in very shallow water environments.<ref>{{Cite journal |last1=Ivantsov |first1=Andrey |last2=Zakrevskaya |first2=Maria |date=July 2022 |title=Dickinsonia : mobile and adhered |url=https://www.cambridge.org/core/product/identifier/S0016756821000194/type/journal_article |journal=Geological Magazine |language=en |volume=159 |issue=7 |pages=1118–1133 |doi=10.1017/S0016756821000194 |issn=0016-7568}}</ref>
 == Discovery ==
@@ Line 55: / Line 54: @@
 === Taphonomy ===
-As a rule, ''Dickinsonia'' fossils are preserved as negative impressions ("death masks") on the bases of sandstone beds. Such fossils are imprints of the upper sides of the benthic organisms that have been buried under the sand.<ref name="Ivantsov2009">{{cite journal |last=Ivantsov |first=A.Y. |year=2009 |title=A new reconstruction of ''Kimberella'', a problematic Vendian Metazoan |journal=Paleontological Journal |volume=43 |issue=6 |pages=601–611 |doi=10.1134/S003103010906001X |s2cid=85676210}}</ref><ref name="Gehling1999">{{Cite journal |last=Gehling |first=J.G. |year=1999 |title=Microbial mats in terminal Proterozoic siliciclastics; Ediacaran death masks |url=http://palaios.geoscienceworld.org/cgi/content/abstract/14/1/40 |journal=PALAIOS |volume=14 |issue=1 |pages=40–57 |bibcode=1999Palai..14...40G |doi=10.2307/3515360 |jstor=3515360}}</ref> The imprints formed as a result of cementation of the sand before complete decomposition of the body. The mechanism of cementation is not quite clear; among many possibilities, the process could have arisen from conditions which gave rise to pyrite "death masks"<ref name="Gehling1999" /> on the decaying body, or perhaps it was due to the carbonate cementation of the sand.<ref name="Serezhnikova2011">{{cite book |last=Serezhnikova Z |first=E.A. |title=Advances in Stromatolite Geobiology |year=2011 |isbn=978-3-642-10414-5 |series=Lecture Notes in Earth Sciences |volume=131 |pages=525–535 |chapter=Microbial Binding as a Probable Cause of Taphonomic Variability of Vendian Fossils: Carbonate Casting? |doi=10.1007/978-3-642-10415-2_31}}</ref> The imprints of the bodies of organisms are often strongly compressed, distorted, and sometimes partly extend into the overlying rock. These deformations appear to show attempts by the organisms to escape from the falling sediment.<ref name="Ivantsov20113"/><ref name="Ivantsov20132"/><ref name="Runnegar1982">{{cite journal |last=Runnegar |first=Bruce |year=1982 |title=Oxygen requirements, biology and phylogenetic significance of the late Precambrian worm ''Dickinsonia'', and the evolution of the burrowing habit |journal=Alcheringa: An Australasian Journal of Palaeontology |volume=6 |issue=3 |pages=223–239 |doi=10.1080/03115518208565415}}</ref>
+As a rule, ''Dickinsonia'' fossils are preserved as negative impressions ("death masks") on the bases of sandstone beds. Such fossils are imprints of the upper sides of the benthic organisms that have been buried under the sand.<ref name=Ivantsov2009>{{cite journal |last=Ivantsov |first=A.Y. |year=2009 |title=A new reconstruction of ''Kimberella'', a problematic Vendian Metazoan |journal=Paleontological Journal |volume=43 |issue=6 |pages=601–611 |doi=10.1134/S003103010906001X |s2cid=85676210}}</ref><ref name="Gehling1999">{{Cite journal |last=Gehling |first=J.G. |year=1999 |title=Microbial mats in terminal Proterozoic siliciclastics; Ediacaran death masks |url=http://palaios.geoscienceworld.org/cgi/content/abstract/14/1/40 |journal=PALAIOS |volume=14 |issue=1 |pages=40–57 |bibcode=1999Palai..14...40G |doi=10.2307/3515360 |jstor=3515360}}</ref> The imprints formed as a result of cementation of the sand before complete decomposition of the body. The mechanism of cementation is not quite clear; among many possibilities, the process could have arisen from conditions which gave rise to pyrite "death masks"<ref name="Gehling1999" /> on the decaying body, or perhaps it was due to the carbonate cementation of the sand.<ref name="Serezhnikova2011">{{cite book |last=Serezhnikova Z |first=E.A. |title=Advances in Stromatolite Geobiology |year=2011 |isbn=978-3-642-10414-5 |series=Lecture Notes in Earth Sciences |volume=131 |pages=525–535 |chapter=Microbial Binding as a Probable Cause of Taphonomic Variability of Vendian Fossils: Carbonate Casting? |doi=10.1007/978-3-642-10415-2_31}}</ref> The imprints of the bodies of organisms are often strongly compressed, distorted, and sometimes partly extend into the overlying rock. These deformations appear to show attempts by the organisms to escape from the falling sediment.<ref name=Ivantsov2011/><ref name=Ivantsov2013/><ref name=Runnegar1982>{{cite journal |last=Runnegar |first=Bruce |year=1982 |title=Oxygen requirements, biology and phylogenetic significance of the late Precambrian worm ''Dickinsonia'', and the evolution of the burrowing habit |journal=Alcheringa: An Australasian Journal of Palaeontology |volume=6 |issue=3 |pages=223–239 |doi=10.1080/03115518208565415}}</ref>
 Rarely, ''Dickinsonia'' have been preserved as a cast in massive sandstone lenses, where it occurs together with ''[[Pteridinium]]'', ''[[Rangea]]'' and some others.<ref name="Grazhdankin20042">{{cite journal |last=Grazhdankin |first=Dima |year=2004 |title=Patterns of distribution in the Ediacaran biotas: facies versus biogeography and evolution |url=http://paleobiol.geoscienceworld.org/cgi/reprint/30/2/203.pdf |journal=Paleobiology |volume=30 |issue=2 |pages=203–221 |doi=10.1666/0094-8373(2004)030<0203:PODITE>2.0.CO;2 |s2cid=129376371}}</ref><ref name="Keller_Fedonkin_1976">{{cite journal |last1=Keller |first1=B.M. |last2=Fedonkin |first2=M.A. |year=1976 |title=New records of fossils in the Valdaian group of the precambrian on the Syuz'ma River |url=http://vend.paleo.ru/pub/Keller_Fedonkin_1976.pdf |url-status=dead |journal=Izvestiya Akademii Nauk SSR |series=Seriya Geologicheskaya |language=ru |volume=3 |pages=38–44 |archive-url=https://web.archive.org/web/20070927022652/http://vend.paleo.ru/pub/Keller_Fedonkin_1976.pdf |archive-date=2007-09-27}}</ref><ref name="Keller_Fedonkin_1977">{{cite journal |last1=Keller |first1=B. M. |last2=Fedonkin |first2=M. A. |year=1977 |title=New organic fossil finds in the Precambrian Valday series along the Syuz'ma River |journal=International Geology Review |volume=19 |issue=8 |pages=924–930 |bibcode=1977IGRv...19..924K |doi=10.1080/00206817709471091}}</ref><ref name="Gehling_Droser2013">{{cite journal |last1=Gehling |first1=J.G. |last2=Droser |first2=M.L. |year=2013 |title=How well do fossil assemblages of the Ediacara Biota tell time? |journal=Geology |volume=41 |issue=4 |pages=447–450 |bibcode=2013Geo....41..447G |doi=10.1130/G33881.1}}</ref> These specimens are products of events where organisms were first stripped from the sea-floor, transported and deposited within sand flow.<ref name="Grazhdankin20042" /><ref name="Gehling_Droser2013" /> In such cases, stretched and ripped ''Dickinsonia'' occur. The first such specimen was described as a separate genus and species, ''Chondroplon bilobatum''<ref name="Wade1971">{{cite journal |last=Wade |first=M. |year=1971 |title=Bilateral Precambrian Chondrophores from the Ediacara Fauna, South Australia |journal=Proceedings of the Royal Society of Victoria |volume=84 |issue=1 |pages=183–188}}</ref> and later re-identified as ''Dickinsonia''.
@@ Line 62: / Line 61: @@
 === Species ===
-Since 1947, a total of nine species have been described, of which three are currently considered valid:<ref name="Evans2023">{{cite journal |last1=Evans |first1=S.D. |last2=Hunt |first2=G. |last3=Gehling |first3=J.G. |last4=Sperling |first4=E. A. |last5=Droser |first5=M.L. |year=2023 |title=Species of Dickinsonia Sprigg from the Ediacaran of South Australia |journal=Palaeontology |volume=66 |issue=e12635 |pages=1–21 |doi=10.1111/pala.12635 |doi-access=free}}</ref>
+Since 1947, a total of nine species have been described, of which three are currently considered valid:<ref name=Evans2023>{{cite journal |last1=Evans |first1=S.D. |last2=Hunt |first2=G. |last3=Gehling |first3=J.G. |last4=Sperling |first4=E. A. |last5=Droser |first5=M.L. |year=2023 |title=Species of ''Dickinsonia sprigg'' from the Ediacaran of South Australia |journal=Palaeontology |volume=66 |issue=1–21 |pages=e12635 |doi=10.1111/pala.12635 |doi-access=free}}</ref>
-{| class="wikitable sortable"
+:{| class="wikitable sortable"
-!Species
+! Species
-!Authority
+! Authority
-!Location
+! Location
-!Status
+! Status
-!Notes
+! Notes
+! Refs
 |-
+| ''Dickinsonia brachina''
-|''Dickinsonia brachina'' <ref name="Wade1972">{{cite journal |last=Wade |first=M. |year=1972 |title=''Dickinsonia'': Polychaete worms from the late Precambrian Ediacara fauna, South Australia |journal=Mem. Queensl. Mus. |volume=16 |issue=2 |pages=171–190}}</ref>
-|Wade, 1972
+| {{harvp|Wade|1972}}
-|Australia
+| Australia
-|invalid
+| invalid
-|Synonym of ''Dickinsonia tenuis''
+| synonym of ''D.&nbsp;tenuis''
+| <ref name=Wade1972>{{cite journal |last=Wade |first=M. |year=1972 |title=''Dickinsonia'': Polychaete worms from the late Precambrian Ediacara fauna, South Australia |journal=Mem. Queensl. Mus. |volume=16 |issue=2 |pages=171–190 }}</ref>
 |-
+| ''Dickinsonia costata''
-|''Dickinsonia costata'' <ref name="Sprigg19472">{{cite journal |last=Sprigg |first=Reg C. |year=1947 |title=Early Cambrian (?) jellyfishes from the Flinders Ranges, South Australia |url=http://www.samuseum.sa.gov.au/Journals/TRSSA/TRSSA_V071/trssa_v071_p212p224.pdf |url-status=dead |journal=Trans. R. Soc. S. Aust. |volume=71 |pages=212–24 |archive-url=https://web.archive.org/web/20070929092905/http://www.samuseum.sa.gov.au/Journals/TRSSA/TRSSA_v071/trssa_v071_p212p224.pdf |archive-date=2007-09-29}}</ref>
+| {{harvp|Sprigg|1947a}}
-|Sprigg, 1947
-|Australia, Russia and Ukraine
+| Australia, Russia, and Ukraine
-|valid
+| valid
-|Unlike other species, ''D. costata'' has comparatively rounded body and fewer, wider segments / isomers.
+| {{efn|group=upper-alpha|Unlike other species, ''D.&nbsp;costata'' has comparatively rounded body and fewer, wider segments / isomers.}}
+| <ref name=Sprigg19472>{{cite journal |last=Sprigg |first=Reg C. |year=1947a |title=Early Cambrian (?) jellyfishes from the Flinders Ranges, South Australia |journal=Trans. R. Soc. S. Aust. |volume=71 |pages=212–24 |url=http://www.samuseum.sa.gov.au/Journals/TRSSA/TRSSA_V071/trssa_v071_p212p224.pdf |url-status=dead |archive-url=https://web.archive.org/web/20070929092905/http://www.samuseum.sa.gov.au/Journals/TRSSA/TRSSA_v071/trssa_v071_p212p224.pdf |archive-date=2007-09-29}}</ref>
 |-
+| ''Dickinsonia elongata''
-|''Dickinsonia elongata'' <ref name="Glaessner1966">{{cite journal |last1=Glaessner |first1=M.F. |last2=Wade |first2=M. |year=1966 |title=The late Precambrian fossils from Ediacara, South Australia |url=http://palaeontology.palass-pubs.org/pdf/Vol%209/Pages%20599-628.pdf |journal=Palaeontology |volume=9 |issue=4 |pages=599}}</ref>
-|Glaessner & Wade, 1966
+| {{harvp|Glaessner|Wade|1966}}
-|Australia
+| Australia
-|invalid
+| invalid
-|Synonym of ''Dickinsonia costata''
+| synonym of ''D.&nbsp;costata''
+| <ref name=Glaessner1966>{{cite journal |last1=Glaessner |first1=M.F. |last2=Wade |first2=M. |year=1966 |title=The late Precambrian fossils from Ediacara, South Australia |url=http://palaeontology.palass-pubs.org/pdf/Vol%209/Pages%20599-628.pdf |journal=Palaeontology |volume=9 |issue=4 |pages=599}}</ref>
 |-
-|''Dickinsonia lissa''<ref name="Wade1972" />
+| ''Dickinsonia lissa''
-|Wade, 1972
+| {{harvp|Wade|1972}}
-|Australia
+| Australia
-|invalid
+| invalid
-|Synonym of ''Dickinsonia tenuis''
+| synonym of ''D.&nbsp;tenuis''
+| <ref name=Wade1972/>
 |-
+| {{nobr|''Dickinsonia menneri'' &thinsp;}}
-|''Dickinsonia menneri''<ref name="Keller_Fedonkin_19762">{{cite journal |last1=Keller |first1=B.M. |last2=Fedonkin |first2=M.A. |year=1976 |title=New records of fossils in the Valdaian group of the precambrian on the Syuz'ma River |url=http://vend.paleo.ru/pub/Keller_Fedonkin_1976.pdf |url-status=dead |journal=Izvestiya Akademii Nauk SSR |series=Seriya Geologicheskaya |language=ru |volume=3 |pages=38–44 |archive-url=https://web.archive.org/web/20070927022652/http://vend.paleo.ru/pub/Keller_Fedonkin_1976.pdf |archive-date=2007-09-27}}</ref>
-|Keller, 1976
+| {{harvp|Keller|1976}}
-|Russia
+| Russia
-|valid
+| valid
-|''D.&nbsp;menneri'' is a small organism up to 8&nbsp;mm in length, and strongly resembles juvenile specimens of ''D.&nbsp;costata'' with its small number of isomers and well-marked head. ''D.&nbsp;menneri'' differs from juvenile ''D.&nbsp;costata'' by its slightly more elongated form.
+| {{efn|group=upper-alpha|''Dickinsonia menneri'' originally was identified as ''[[Vendomia]]'' but re-classified as ''Dickinsonia'' by {{harvp|Ivantsov|2007}}<ref name=Ivantsov2007/>}}{{efn|group=upper-alpha|''D.&nbsp;menneri'' is a small organism up to 8&nbsp;mm in length, and strongly resembles juvenile specimens of ''D.&nbsp;costata'' with its small number of isomers and well-marked head. ''D.&nbsp;menneri'' differs from juvenile ''D.&nbsp;costata'' by its slightly more elongated form.}}
+| <ref name=Keller_Fedonkin_19762>{{cite journal |last1=Keller |first1=B.M. |last2=Fedonkin |first2=M.A. |year=1976 |title=New records of fossils in the Valdaian group of the precambrian on the Syuz'ma River |journal=Izvestiya Akademii Nauk SSR |series=Seriya Geologicheskaya |language=ru |volume=3 |pages=38–44 |url=http://vend.paleo.ru/pub/Keller_Fedonkin_1976.pdf |url-status=dead |archive-url=https://web.archive.org/web/20070927022652/http://vend.paleo.ru/pub/Keller_Fedonkin_1976.pdf |archive-date=2007-09-27}}</ref>
-Originally classified as ''Vendomia'', it was re-identified as ''Dickinsonia'' by Ivantsov (2007)<ref name="Ivantsov20072">{{cite journal |last=Ivantsov |first=A. Y. |year=2007 |title=Small Vendian transversely articulated fossils |url=https://www.academia.edu/2352394 |journal=Paleontological Journal |volume=41 |issue=2 |pages=113–122 |doi=10.1134/S0031030107020013 |s2cid=86636748}}</ref>
 |-
+| ''Dickinsonia minima''
-|''Dickinsonia minima'' <ref name="Sprigg1949">{{cite journal |last=Sprigg |first=R.C. |year=1949 |title=Early Cambrian "jellyfishes" of Ediacara, South Australia, and Mount John, Kimberley District, Western Australia |url=http://www.samuseum.sa.gov.au/Journals/TRSSA/TRSSA_V073/trssa_v073_p72p99.pdf |journal=Transactions of the Royal Society of South Australia |volume=73 |pages=72–99}}{{dead link|date=September 2017|bot=InternetArchiveBot|fix-attempted=yes}}</ref>
-|Sprigg, 1949
+| {{harvp|Sprigg|1949}}
-|Australia
+| Australia
-|invalid
+| invalid
-|Synonym of ''Dickinsonia costata''
+| synonym of ''D.&nbsp;costata''
+| <ref name=Sprigg1949>{{cite journal |last=Sprigg |first=R.C. |year=1949 |title=Early Cambrian "jellyfishes" of Ediacara, South Australia, and Mount John, Kimberley District, Western Australia |journal=Transactions of the Royal Society of South Australia |volume=73 |pages=72–99 |url=http://www.samuseum.sa.gov.au/Journals/TRSSA/TRSSA_V073/trssa_v073_p72p99.pdf |url-status=dead }}{{dead link|date=September 2017|bot=InternetArchiveBot|fix-attempted=yes}}</ref>
 |-
+| ''Dickinsonia rex''
-|''Dickinsonia rex'' <ref name="Jenkins1992">{{Cite book |last=Jenkins |first=R.J.F. |title=Origin and early evolution of the Metazoa |publisher=Springer |year=1992 |isbn=978-0-306-44067-0 |editor-last1=Lipps |editor-first1=J. |location=New York, NY |pages=131–176 |chapter=Functional and ecological aspects of Ediacarian assemblages |oclc=231467647 |editor-last2=Signor |editor2-first=P.W. |chapter-url=https://books.google.com/books?id=gUQMKiJOj64C&pg=PA165}}</ref>
-|Jenkins, 1992
+| {{harvp|Jenkins|1992}}
-|Australia
+| Australia
-|invalid
+| invalid
-|Synonym of ''Dickinsonia tenuis''
+| synonym of ''D.&nbsp;tenuis''
+| <ref name=Jenkins1992>{{cite book |last=Jenkins |first=R.J.F. |year=1992 |chapter=Functional and ecological aspects of Ediacarian assemblages |editor1-last=Lipps |editor1-first=J. |editor2-last=Signor |editor2-first=P.W. |title=Origin and Early Evolution of the Metazoa |publisher=Springer |isbn=978-0-306-44067-0 |location=New York, NY |pages=131–176 |oclc=231467647 |chapter-url=https://books.google.com/books?id=gUQMKiJOj64C&pg=PA165}}</ref>
 |-
+| ''Dickinsonia spriggi''
-|''Dickinsonia spriggi'' <ref name="Harrington_Moore1955">{{cite journal |last1=Harrington |first1=N.J. |last2=Moore |first2=R.C. |year=1955 |title=Kansas Pennsylvanian and other jellyfishes |url=http://www.kgs.ku.edu/Publications/Bulletins/114_5/index.html |journal=Bull. Kansas Geol. Surv. |volume=114 |issue=5 |pages=153–163}}</ref>
-|Harrington & Moore, 1955
+| {{harvp|Harrington|Moore|1955}}
-|Australia
+| Australia
-|invalid
+| invalid
-|Synonym of ''Dickinsonia costata''
+| synonym of ''D.&nbsp;costata''
+| <ref name=Harrington_Moore1955>{{cite journal |last1=Harrington |first1=N.J. |last2=Moore |first2=R.C. |year=1955 |title=Kansas Pennsylvanian and other jellyfishes |journal=Bull. Kansas Geol. Surv. |volume=114 |issue=5 |pages=153–163 |url=http://www.kgs.ku.edu/Publications/Bulletins/114_5/index.html}}</ref>
 |-
-|''Dickinsonia tenuis'' <ref name="Glaessner1966" />
+| ''Dickinsonia tenuis''
-|Glaessner & Wade, 1966
+| {{harvp|Glaessner|Wade|1966}}
-|Australia and Russia
+| Australia and Russia
-|valid
+| valid
-|Strongly resembles ''D.&nbsp;costata'', but differs from it by more narrow and numerous segments, sparingly lengthened oval form of the body.
+| {{efn|group=upper-alpha|''Dickinsonia tenuis'' strongly resembles ''D.&nbsp;costata'', but differs from it by more narrow and numerous segments, sparingly lengthened oval form of the body.}}
+| <ref name=Glaessner1966/>
 |}
+::{{notelist|group=upper-alpha}}
-A claimed specimen of ''Dickinsonia'' from India was later determined to be the remains of a beehive.<ref>{{Cite journal |last1=Pandey |first1=S. K. |last2=Ahmad |first2=Shamim |last3=Sharma |first3=Mukund |date=2023-03-09 |title=Dickinsonia tenuis reported by Retallack et al. 2021 is not a fossil, instead an impression of an extant 'fallen beehive' |url=https://link.springer.com/10.1007/s12594-023-2312-2 |journal=Journal of the Geological Society of India |language=en |volume=99 |issue=3 |pages=311–316 |doi=10.1007/s12594-023-2312-2 |issn=0974-6889}}</ref>
+A claimed specimen of ''Dickinsonia'' from India was later determined to be the remains of a beehive.<ref>{{cite journal |last1=Pandey |first1=S.K. |last2=Ahmad |first2=Shamim |last3=Sharma |first3=Mukund |date=2023-03-09 |title=Dickinsonia tenuis reported by Retallack ''et al.'' 2021 is not a fossil, instead an impression of an extant 'fallen beehive' |journal=Journal of the Geological Society of India |lang=en |volume=99 |issue=3 |pages=311–316 |doi=10.1007/s12594-023-2312-2 |issn=0974-6889 |url=https://link.springer.com/10.1007/s12594-023-2312-2 }}</ref>
 === External relationships ===
-''Dickinsonia'' is classified as part of the group [[Proarticulata]] or [[Dickinsoniomorpha]].<ref name=":3" /> Proarticulata includes a number of morphologically similar organisms, such ''[[Spriggina]]'', ''[[Yorgia]]'', ''[[Andiva]]'' and ''[[Cephalonega]],'' which share the same segmented articulation.<ref name="Ivantsov_20192">{{Cite journal |author1=A.Y. Ivantsov |author2=M.A. Fedonkin |author3=A.L. Nagovitsyn |author4=M.A. Zakrevskaya |year=2019 |title=''Cephalonega'', a new generic name, and the system of Vendian Proarticulata |url=https://www.researchgate.net/publication/335988735 |journal=Paleontological Journal |volume=53 |issue=5 |pages=447–454 |doi=10.1134/S0031030119050046 |s2cid=203853224}}</ref> The affinities of Proarticulata to other organisms, including to other members of the [[Ediacaran biota]], like [[rangeomorph]]s, have long been contentious.''<ref name="Bobrovskiy" />'' It has been historically proposed that most Ediacaran organisms were closely related to each other, as part of the grouping "[[Vendobionta]]",<ref name="Seilacher1992" /> though recent authors argue that this grouping as a whole is likely to be [[Polyphyly|polyphyletic]].''<ref name="Bobrovskiy" />'' [[Gregory Retallack]] has proposed that the fossils of ''Dickinsonia'' and other Ediacaran biota represent lichens that grew in a terrestrial environment,<ref>{{Cite journal |last=Retallack |first=Gregory J. |date=January 2013 |title=Ediacaran life on land |url=https://www.nature.com/articles/nature11777 |journal=Nature |language=en |volume=493 |issue=7430 |pages=89–92 |doi=10.1038/nature11777 |issn=0028-0836}}</ref> but this has been broadly rejected by other authors, who argue that a marine environment of deposition better fits available evidence.<ref>{{Cite journal |last1=Xiao |first1=Shuhai |last2=Knauth |first2=L. Paul |date=January 2013 |title=Fossils come in to land |journal=Nature |language=en |volume=493 |issue=7430 |pages=28–29 |doi=10.1038/nature11765 |issn=0028-0836|doi-access=free }}</ref>''<ref name="Bobrovskiy" />''<ref>{{Cite journal |last1=Xiao |first1=Shuhai |last2=Droser |first2=Mary |last3=Gehling |first3=James G. |last4=Hughes |first4=Ian V. |last5=Wan |first5=Bin |last6=Chen |first6=Zhe |last7=Yuan |first7=Xunlai |date=March 2014 |title=Affirming life aquatic for the Ediacara biota in China and Australia: REPLY |url=http://pubs.geoscienceworld.org/geology/article/42/3/e326/131566/Affirming-life-aquatic-for-the-Ediacara-biota-in |journal=Geology |language=en |volume=42 |issue=3 |pages=e326 |doi=10.1130/G35364Y.1 |issn=1943-2682}}</ref> Other proposal have included giant [[protist]]s, as proposed by [[Adolf Seilacher]].<ref>{{Cite journal |last=Seilacher |first=A. |date=January 2007 |title=The nature of vendobionts |url=https://www.lyellcollection.org/doi/10.1144/SP286.28 |journal=Geological Society, London, Special Publications |language=en |volume=286 |issue=1 |pages=387–397 |doi=10.1144/SP286.28 |issn=0305-8719}}</ref> Most modern research suggest that ''Dickinsonia'' and other proarticulatans are likely to be animals, possibly belonging to [[Eumetazoa]].<ref name=":4" /><ref name=":1" /><ref name=":3" /> A chemical study of Russian specimens found that they were enriched with [[cholesterol]], which is only produced by animals, supporting an animal affinity,''<ref name="Bobrovskiy" />'' though these results have been questioned by other authors, who consider the association between the cholesterol molecules and the ''Dickinsonia'' fossils to not be definitive.<ref name="Love2021" /> Within Animalia, a number of affinities have been proposed, including as stem-eumetazoans forming a clade with rangeomorphs,<ref>{{Cite journal |last1=Hoyal Cuthill |first1=Jennifer F. |last2=Han |first2=Jian |date=November 2018 |editor-last=Álvaro |editor-first=Javier |title=Cambrian petalonamid Stromatoveris phylogenetically links Ediacaran biota to later animals |journal=Palaeontology |language=en |volume=61 |issue=6 |pages=813–823 |doi=10.1111/pala.12393 |issn=0031-0239|doi-access=free }}</ref> to [[Placozoa]],<ref name="Sperling2008">{{cite journal |author1=Sperling, Erik |display-authors=etal |year=2008 |title=A placozoan affinity for ''Dickinsonia'' and the evolution of late Precambrian metazoan feeding modes |url=http://gsa.confex.com/gsa/2008AM/finalprogram/abstract_151743.htm |url-status=dead |journal=Geological Society of America |series=Abstracts with Programs |volume=40 |issue=6 |pages=508 |archive-url=https://web.archive.org/web/20180228161702/https://gsa.confex.com/gsa/2008AM/finalprogram/abstract_151743.htm |archive-date=2018-02-28 |access-date=2008-10-27}}</ref> and to [[Cnidaria]].<ref>J.W. Valentine ''Dickinsonia'' as a polypoid organism Paleobiology, 18 (1992), pp. 378-382</ref> A number of researchers have proposed close affinities to [[Bilateria]], based on the bilateral or nearly bilateral organisation of proarticulatans,<ref name=":3" /><ref name=":0" /> though proarticulatans are not likely to be a member of the bilaterian [[crown group]].<ref name=":1" />
+''Dickinsonia'' is classified as part of the group [[Proarticulata]] or [[Dickinsoniomorpha]].<ref name=":3" /> Proarticulata includes a number of morphologically similar organisms, such ''[[Spriggina]]'', ''[[Yorgia]]'', ''[[Andiva]]'' and ''[[Cephalonega]]'', which share the same segmented articulation.<ref name=Ivantsov_2019>{{Cite journal |first1=A.Y. |last1=Ivantsov |first2=M.A. |last2=Fedonkin |first3=A.L. |last3=Nagovitsyn |author4=M.A. Zakrevskaya |year=2019 |title=''Cephalonega'', a new generic name, and the system of Vendian Proarticulata |url=https://www.researchgate.net/publication/335988735 |journal=Paleontological Journal |volume=53 |issue=5 |pages=447–454 |doi=10.1134/S0031030119050046 |s2cid=203853224 }}</ref> The affinities of Proarticulata to other organisms, including to other members of the [[Ediacaran biota]], like [[rangeomorph]]s, have long been contentious.<ref name=Bobrovskiy/> It has been historically proposed that most Ediacaran organisms were closely related to each other, as part of the grouping "[[Vendobionta]]",<ref name=Seilacher1992/> though recent authors argue that this grouping as a whole is likely to be [[Polyphyly|polyphyletic]].<ref name=Bobrovskiy/> [[Gregory Retallack]] has proposed that the fossils of ''Dickinsonia'' and other Ediacaran biota represent lichens that grew in a terrestrial environment,<ref>{{cite journal |last=Retallack |first=Gregory J. |date=January 2013 |title=Ediacaran life on land |journal=Nature |lang=en |volume=493 |issue=7430 |pages=89–92 |doi=10.1038/nature11777 |issn=0028-0836 |url=https://www.nature.com/articles/nature11777}}</ref> but this has been broadly rejected by other authors, who argue that a marine environment of deposition better fits available evidence.<ref>{{cite journal |last1=Xiao |first1=Shuhai |last2=Knauth |first2=L. Paul |date=January 2013 |title=Fossils come in to land |journal=Nature |lang=en |volume=493 |issue=7430 |pages=28–29 |doi=10.1038/nature11765 |issn=0028-0836|doi-access=free }}</ref><ref name=Bobrovskiy/><ref>{{Cite journal |last1=Xiao |first1=Shuhai |last2=Droser |first2=Mary |last3=Gehling |first3=James G. |last4=Hughes |first4=Ian V. |last5=Wan |first5=Bin |last6=Chen |first6=Zhe |last7=Yuan |first7=Xunlai |date=March 2014 |title=Affirming life aquatic for the Ediacara biota in China and Australia: REPLY |url=http://pubs.geoscienceworld.org/geology/article/42/3/e326/131566/Affirming-life-aquatic-for-the-Ediacara-biota-in |journal=Geology |language=en |volume=42 |issue=3 |pages=e326 |doi=10.1130/G35364Y.1 |issn=1943-2682}}</ref> Other proposal have included giant [[protist]]s, as proposed by [[Adolf Seilacher]].<ref>{{Cite journal |last=Seilacher |first=A. |date=January 2007 |title=The nature of vendobionts |url=https://www.lyellcollection.org/doi/10.1144/SP286.28 |journal=Geological Society, London, Special Publications |language=en |volume=286 |issue=1 |pages=387–397 |doi=10.1144/SP286.28 |issn=0305-8719}}</ref> Most modern research suggest that ''Dickinsonia'' and other proarticulatans are likely to be animals, possibly belonging to [[Eumetazoa]].<ref name=":4"/><ref name=":1"/><ref name=":3"/> A chemical study of Russian specimens found that they were enriched with [[cholesterol]], which is only produced by animals, supporting an animal affinity,<ref name=Bobrovskiy/> though these results have been questioned by other authors, who consider the association between the cholesterol molecules and the ''Dickinsonia'' fossils to not be definitive.<ref name=Love2021/> Within Animalia, a number of affinities have been proposed, including as stem-eumetazoans forming a clade with rangeomorphs,<ref>{{cite journal |last1=Hoyal Cuthill |first1=Jennifer F. |last2=Han |first2=Jian |date=November 2018 |editor-last=Álvaro |editor-first=Javier |title=Cambrian petalonamid Stromatoveris phylogenetically links Ediacaran biota to later animals |journal=Palaeontology |language=en |volume=61 |issue=6 |pages=813–823 |doi=10.1111/pala.12393 |issn=0031-0239|doi-access=free }}</ref> to [[Placozoa]],<ref name="Sperling2008">{{cite journal |author1=Sperling, Erik |display-authors=etal |year=2008 |title=A placozoan affinity for ''Dickinsonia'' and the evolution of late Precambrian metazoan feeding modes |url=http://gsa.confex.com/gsa/2008AM/finalprogram/abstract_151743.htm |url-status=dead |journal=Geological Society of America |series=Abstracts with Programs |volume=40 |issue=6 |pages=508 |archive-url=https://web.archive.org/web/20180228161702/https://gsa.confex.com/gsa/2008AM/finalprogram/abstract_151743.htm |archive-date=2018-02-28 |access-date=2008-10-27}}</ref> and to [[Cnidaria]].<ref>J.W. Valentine ''Dickinsonia'' as a polypoid organism Paleobiology, 18 (1992), pp. 378-382</ref> A number of researchers have proposed close affinities to [[Bilateria]], based on the bilateral or nearly bilateral organisation of proarticulatans,<ref name=":3" /><ref name=":0" /> though proarticulatans are not likely to be a member of the bilaterian [[crown group]].<ref name=":1" />
 ==References==
-{{reflist}}
+{{reflist|25em}}
 {{Portal|Paleontology}}