Biology:Dickinsonia

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Short description: Extinct genus of early animals

Dickinsonia
Temporal range: Late Ediacaran, 567–550 Ma
DickinsoniaCostata.jpg
Cast of Dickinsonia costata from Australia
Scientific classification e
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

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] The discovery of cholesterol molecules in fossils of Dickinsonia lends support to the idea that Dickinsonia was an animal,[7] though these results have been questioned.[8]

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.[9] 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][10] The body is divided into two by a midline ridge or groove,[2][10][11] except for a single unpaired segment at one end, dubbed the "anterior most unit" suggested to represent the front of the organism.[11] Is it disputed whether the segments are offset from each other following glide reflection, and are thus isomers,[2][10][12][13] 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.[11][14] The number of segments/isomer pairs varies from 12 in smaller individuals to 74 in the largest Australian specimens.[14]

The body of Dickinsonia is suggested to have been sack-like, with the outer layer being made of a resistant but unmineralised material.[13] Some specimens from Russia show the presence of branched internal structures.[15][13] Some authors have suggested that the underside of the body bore cilia, as well as infolded pockets.[13]

Dickinsonia is suggested to have grown by adding a new pair of segments/isomers at the end opposite the unpaired "anterior most unit".[11][16] Dickinsonia probably exhibited indeterminate growth (having no maximum size), though it is suggested that the addition of new segments slowed down later in growth.[17] Deformed specimens from Russia indicate that individuals of Dickinsonia could regenerate after being damaged.[16]

  • 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.[13] These trace fossils have been assigned to the genus Epibaion.[12][18][19] 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.[20]

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,[21] described Dickinsonia, naming it after Ben Dickinson, then Director of Mines for South Australia, and head of the government department that employed Sprigg.[22] Additional specimens of Dickinsonia are also known from the Mogilev Formation in the Dniester River Basin of Podolia, Ukraine ,[23] the Lyamtsa, Verkhovka, Zimnegory and Yorga Formations in the White Sea area of the Arkhangelsk Region, Chernokamen Formation of the Central Urals, Russia ,[9] (these deposits have been dated to 567–550 Myr.[24][25][26]), the Dengying Formation in the Yangtze Gorges area, South China. (ca. 551–543 Ma).[27]

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.[28][29] 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"[29] on the decaying body, or perhaps it was due to the carbonate cementation of the sand.[30] 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.[12][18][31]

Rarely, Dickinsonia have been preserved as a cast in massive sandstone lenses, where it occurs together with Pteridinium, Rangea and some others.[32][33][34][35] These specimens are products of events where organisms were first stripped from the sea-floor, transported and deposited within sand flow.[32][35] In such cases, stretched and ripped Dickinsonia occur. The first such specimen was described as a separate genus and species, Chondroplon bilobatum[36] 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:[37]

Species Authority Location Status Notes
Dickinsonia brachina [38] Wade, 1972 Australia invalid Synonym of Dickinsonia tenuis
Dickinsonia costata [39] Sprigg, 1947 Australia, Russia and Ukraine valid Unlike other species, D. costata has comparatively rounded body and fewer, wider segments / isomers.
Dickinsonia elongata [40] Glaessner & Wade, 1966 Australia invalid Synonym of Dickinsonia costata
Dickinsonia lissa[38] Wade, 1972 Australia invalid Synonym of Dickinsonia tenuis
Dickinsonia menneri[41] Keller, 1976 Russia valid 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.

Originally classified as Vendomia, it was re-identified as Dickinsonia by Ivantsov (2007)[2]

Dickinsonia minima [42] Sprigg, 1949 Australia invalid Synonym of Dickinsonia costata
Dickinsonia rex [43] Jenkins, 1992 Australia invalid Synonym of Dickinsonia tenuis
Dickinsonia spriggi [44] Harrington & Moore, 1955 Australia invalid Synonym of Dickinsonia costata
Dickinsonia tenuis [40] Glaessner & Wade, 1966 Australia and Russia valid 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.[45]

External relationships

Dickinsonia is classified as part of the group Proarticulata or Dickinsoniomorpha.[13] Proarticulata includes a number of morphologically similar organisms, such Spriggina, Yorgia, Andiva and Cephalonega, which share the same segmented articulation.[46] The affinities of Proarticulata to other organisms, including to other members of the Ediacaran biota, like rangeomorphs, have long been contentious.[7] It has been historically proposed that most Ediacaran organisms were closed 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.[7] Gregory Retallack has proposed that the fossils of Dickinsonia and other Ediacaran biota represent lichens that grew in a terrestrial environment,[47] but this has been broadly rejected by other authors, who argue that a marine environment of deposition better fits available evidence.[48][7][49] Other proposal have included giant protists, as proposed by Adolf Seilacher.[50] Most modern research suggest that Dickinsonia and other proarticulatans are likely to be animals, possibly belonging to Eumetazoa.[17][11][13] A chemical study of Russian specimens found that they were enriched with cholesterol, which is only produced by animals, supporting an animal affinity,[7] 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.[8] Within Animalia, a number of affinities have been proposed, including as stem-eumetazoans forming a clade with rangeomorphs,[51] to Placozoa,[52] and to Cnidaria.[53] A number of reseachers have proposed close affinities to Bilateria, based on the bilateral or nearly bilateral organisation of proarticulatans,[13][3] though proarticulatans are not likely to be a member of the bilaterian crown group.[11]

References

  1. ↑ Hofmann, Hans J. (1988). "An alternative interpretation of the Ediacaran (Precambrian) chondrophore Chondroplon Wade". Alcheringa 12 (4): 315–318. doi:10.1080/03115518808619130. 
  2. ↑ 2.0 2.1 2.2 2.3 2.4 Ivantsov, A. Y. (2007). "Small Vendian transversely articulated fossils". Paleontological Journal 41 (2): 113–122. doi:10.1134/S0031030107020013. https://www.academia.edu/2352394. 
  3. ↑ 3.0 3.1 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. 
  4. ↑ Pflug (1973). "Zur fauna der Nama-Schichten in Südwest-Afrika. IV. Mikroscopische anatomie der petalo-organisme". Palaeontographica (B144): 166–202. 
  5. ↑ 5.0 5.1 Seilacher, Adolf (1992). "Vendobionta and Psammocorallia: lost constructions of Precambrian evolution". Journal of the Geological Society, London 149 (4): 607–613. doi:10.1144/gsjgs.149.4.0607. Bibcode: 1992JGSoc.149..607S. http://jgs.lyellcollection.org/cgi/content/abstract/149/4/607. Retrieved 2007-06-21. 
  6. ↑ McMenamin M. (1998). The Garden of Ediacara. New York: Columbia University Press. ISBN 978-0-231-10559-0. OCLC 228271905.  It lived during the late Ediacaran (part of Precambrian). Dinosaurs a visual encyclopedia. New York: DK Publishing, Inc. 2018-04-03. ISBN 9781465469489. 
  7. ↑ 7.0 7.1 7.2 7.3 7.4 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. doi:10.1126/science.aat7228. PMID 30237355. Bibcode: 2018Sci...361.1246B. 
  8. ↑ 8.0 8.1 Love, G.D.; Zumberge, J.A. (2021). "Emerging patterns in Proterozoic lipid biomarker records". Cambridge Elements 361 (6408). doi:10.1017/9781108847117. ISBN 9781108847117. 
  9. ↑ 9.0 9.1 Fedonkin, 2 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. pp. 326. ISBN 978-0-8018-8679-9. https://books.google.com/books?id=OFKG6SmPNuUC. 
  10. ↑ 10.0 10.1 10.2 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. 
  11. ↑ 11.0 11.1 11.2 11.3 11.4 11.5 Evans, Scott D.; Droser, Mary L.; Gehling, James G. (2017-05-17). Hejnol, Andreas. ed. "Highly regulated growth and development of the Ediacara macrofossil Dickinsonia costata" (in en). PLOS ONE 12 (5): e0176874. doi:10.1371/journal.pone.0176874. ISSN 1932-6203. PMID 28520741. 
  12. ↑ 12.0 12.1 12.2 Ivantsov, A. Y. (2011). "Feeding traces of Proarticulata — the Vendian metazoa". Paleontological Journal 45 (3): 237–248. doi:10.1134/S0031030111030063. https://www.academia.edu/2394670. 
  13. ↑ 13.0 13.1 13.2 13.3 13.4 13.5 13.6 13.7 Ivantsov, Andrey Yu; Zakrevskaya, Maria (2023-02-23). "Body plan of Dickinsonia , the oldest mobile animals" (in en). Earth and Environmental Science Transactions of the Royal Society of Edinburgh 114 (1–2): 95–108. doi:10.1017/S175569102300004X. ISSN 1755-6910. https://www.cambridge.org/core/product/identifier/S175569102300004X/type/journal_article. 
  14. ↑ 14.0 14.1 Dunn, Frances S.; Liu, Alexander G.; Donoghue, Philip C. J. (May 2018). "Ediacaran developmental biology" (in en). Biological Reviews 93 (2): 914–932. doi:10.1111/brv.12379. ISSN 1464-7931. PMID 29105292. 
  15. ↑ Ivantsov, A. Y. (2004). "New Proarticulata from the Vendian of the Arkhangel'sk Region". Paleontological Journal 38 (3): 247–253. http://vend.paleo.ru/pub/Ivantsov_2004_eng.pdf. Retrieved 2007-09-27. 
  16. ↑ 16.0 16.1 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)" (in en). Journal of Paleontology 94 (6): 1019–1033. doi:10.1017/jpa.2020.65. ISSN 0022-3360. https://www.cambridge.org/core/product/identifier/S0022336020000657/type/journal_article. 
  17. ↑ 17.0 17.1 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" (in en). Palaeontology 66 (1). doi:10.1111/pala.12635. ISSN 0031-0239. 
  18. ↑ 18.0 18.1 Ivantsov, A.Y. (2013). "Trace Fossils of Precambrian Metazoans "Vendobionta" and "Mollusks"". Stratigraphy and Geological Correlation 21 (3): 252–264. doi:10.1134/S0869593813030039. Bibcode: 2013SGC....21..252I. https://www.academia.edu/3757185. 
  19. ↑ Ivantsov, A.Y.; Malakhovskaya, Y.E. (2002). "Giant Traces of Vendian Animals". Doklady Earth Sciences 385 (6): 618–622. http://vend.paleo.ru/pub/Ivantsov_et_Malakhovskaya_2002-e.pdf. Retrieved 2008-02-24. 
  20. ↑ Ivantsov, Andrey; Zakrevskaya, Maria (July 2022). "Dickinsonia : mobile and adhered" (in en). Geological Magazine 159 (7): 1118–1133. doi:10.1017/S0016756821000194. ISSN 0016-7568. https://www.cambridge.org/core/product/identifier/S0016756821000194/type/journal_article. 
  21. ↑ Sprigg, Reg C. (1947). "Early Cambrian (?) jellyfishes from the Flinders Ranges, South Australia". Trans. R. Soc. S. Aust. 71: 212–24. http://www.samuseum.sa.gov.au/Journals/TRSSA/TRSSA_V071/trssa_v071_p212p224.pdf. 
  22. ↑ Eig, Karsten. "EDIACARANS: THE FOSSILS THAT SHOULD NOT BE THERE". https://karsteneig.no/2019/05/ediacarans-the-fossils-that-should-not-be-there/. 
  23. ↑ Fedonkin, M. A. (1983). "Non-skeletal fauna of Podoloia, Dniester River valley" (in ru). The Vendian of the Ukraine. Kiev: Naukova Dumka. pp. 128–139. 
  24. ↑ Grazhdankin, Dima (2004). "Patterns of distribution in the Ediacaran biotas: facies versus biogeography and evolution". Paleobiology 30 (2): 203–221. doi:10.1666/0094-8373(2004)030<0203:PODITE>2.0.CO;2. http://paleobiol.geoscienceworld.org/cgi/reprint/30/2/203.pdf. 
  25. ↑ 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. https://www.researchgate.net/publication/327388718. 
  26. ↑ 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. doi:10.1016/j.precamres.2018.08.011. Bibcode: 2018PreR..316..197K. https://www.repository.cam.ac.uk/handle/1810/285052. 
  27. ↑ 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. 
  28. ↑ Ivantsov, A.Y. (2009). "A new reconstruction of Kimberella, a problematic Vendian Metazoan". Paleontological Journal 43 (6): 601–611. doi:10.1134/S003103010906001X. 
  29. ↑ 29.0 29.1 Gehling, J.G. (1999). "Microbial mats in terminal Proterozoic siliciclastics; Ediacaran death masks". PALAIOS 14 (1): 40–57. doi:10.2307/3515360. Bibcode: 1999Palai..14...40G. http://palaios.geoscienceworld.org/cgi/content/abstract/14/1/40. 
  30. ↑ 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. 131. pp. 525–535. doi:10.1007/978-3-642-10415-2_31. ISBN 978-3-642-10414-5. 
  31. ↑ 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. 
  32. ↑ 32.0 32.1 Grazhdankin, Dima (2004). "Patterns of distribution in the Ediacaran biotas: facies versus biogeography and evolution". Paleobiology 30 (2): 203–221. doi:10.1666/0094-8373(2004)030<0203:PODITE>2.0.CO;2. http://paleobiol.geoscienceworld.org/cgi/reprint/30/2/203.pdf. 
  33. ↑ Keller, B.M.; Fedonkin, M.A. (1976). "New records of fossils in the Valdaian group of the precambrian on the Syuz'ma River" (in ru). Izvestiya Akademii Nauk SSR. Seriya Geologicheskaya 3: 38–44. http://vend.paleo.ru/pub/Keller_Fedonkin_1976.pdf. 
  34. ↑ 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. doi:10.1080/00206817709471091. Bibcode: 1977IGRv...19..924K. 
  35. ↑ 35.0 35.1 Gehling, J.G.; Droser, M.L. (2013). "How well do fossil assemblages of the Ediacara Biota tell time?". Geology 41 (4): 447–450. doi:10.1130/G33881.1. Bibcode: 2013Geo....41..447G. 
  36. ↑ Wade, M. (1971). "Bilateral Precambrian Chondrophores from the Ediacara Fauna, South Australia". Proceedings of the Royal Society of Victoria 84 (1): 183–188. 
  37. ↑ 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 (e12635): 1–21. doi:10.1111/pala.12635. 
  38. ↑ 38.0 38.1 Wade, M. (1972). "Dickinsonia: Polychaete worms from the late Precambrian Ediacara fauna, South Australia". Mem. Queensl. Mus. 16 (2): 171–190. 
  39. ↑ Sprigg, Reg C. (1947). "Early Cambrian (?) jellyfishes from the Flinders Ranges, South Australia". Trans. R. Soc. S. Aust. 71: 212–24. http://www.samuseum.sa.gov.au/Journals/TRSSA/TRSSA_V071/trssa_v071_p212p224.pdf. 
  40. ↑ 40.0 40.1 Glaessner, M.F.; Wade, M. (1966). "The late Precambrian fossils from Ediacara, South Australia". Palaeontology 9 (4): 599. http://palaeontology.palass-pubs.org/pdf/Vol%209/Pages%20599-628.pdf. 
  41. ↑ Keller, B.M.; Fedonkin, M.A. (1976). "New records of fossils in the Valdaian group of the precambrian on the Syuz'ma River" (in ru). Izvestiya Akademii Nauk SSR. Seriya Geologicheskaya 3: 38–44. http://vend.paleo.ru/pub/Keller_Fedonkin_1976.pdf. 
  42. ↑ Sprigg, R.C. (1949). "Early Cambrian "jellyfishes" of Ediacara, South Australia, and Mount John, Kimberley District, Western Australia". Transactions of the Royal Society of South Australia 73: 72–99. http://www.samuseum.sa.gov.au/Journals/TRSSA/TRSSA_V073/trssa_v073_p72p99.pdf. [yes|permanent dead link|dead link}}]
  43. ↑ Jenkins, R.J.F. (1992). "Functional and ecological aspects of Ediacarian assemblages". Origin and early evolution of the Metazoa. New York, NY: Springer. pp. 131–176. ISBN 978-0-306-44067-0. OCLC 231467647. https://books.google.com/books?id=gUQMKiJOj64C&pg=PA165. 
  44. ↑ Harrington, N.J.; Moore, R.C. (1955). "Kansas Pennsylvanian and other jellyfishes". Bull. Kansas Geol. Surv. 114 (5): 153–163. http://www.kgs.ku.edu/Publications/Bulletins/114_5/index.html. 
  45. ↑ Pandey, S. K.; Ahmad, Shamim; Sharma, Mukund (2023-03-09). "Dickinsonia tenuis reported by Retallack et al. 2021 is not a fossil, instead an impression of an extant 'fallen beehive'" (in en). Journal of the Geological Society of India 99 (3): 311–316. doi:10.1007/s12594-023-2312-2. ISSN 0974-6889. https://link.springer.com/10.1007/s12594-023-2312-2. 
  46. ↑ A.Y. Ivantsov; M.A. Fedonkin; A.L. Nagovitsyn; 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. https://www.researchgate.net/publication/335988735. 
  47. ↑ Retallack, Gregory J. (January 2013). "Ediacaran life on land" (in en). Nature 493 (7430): 89–92. doi:10.1038/nature11777. ISSN 0028-0836. https://www.nature.com/articles/nature11777. 
  48. ↑ Xiao, Shuhai; Knauth, L. Paul (January 2013). "Fossils come in to land" (in en). Nature 493 (7430): 28–29. doi:10.1038/nature11765. ISSN 0028-0836. 
  49. ↑ 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" (in en). Geology 42 (3): e326. doi:10.1130/G35364Y.1. ISSN 1943-2682. http://pubs.geoscienceworld.org/geology/article/42/3/e326/131566/Affirming-life-aquatic-for-the-Ediacara-biota-in. 
  50. ↑ Seilacher, A. (January 2007). "The nature of vendobionts" (in en). Geological Society, London, Special Publications 286 (1): 387–397. doi:10.1144/SP286.28. ISSN 0305-8719. https://www.lyellcollection.org/doi/10.1144/SP286.28. 
  51. ↑ Hoyal Cuthill, Jennifer F.; Han, Jian (November 2018). Álvaro, Javier. ed. "Cambrian petalonamid Stromatoveris phylogenetically links Ediacaran biota to later animals" (in en). Palaeontology 61 (6): 813–823. doi:10.1111/pala.12393. ISSN 0031-0239. 
  52. ↑ Sperling, Erik (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. http://gsa.confex.com/gsa/2008AM/finalprogram/abstract_151743.htm. Retrieved 2008-10-27. 
  53. ↑ J.W. Valentine Dickinsonia as a polypoid organism Paleobiology, 18 (1992), pp. 378-382

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