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Proarticulata; Temporal range: Late Ediacaran, between 567–550 Ma PreꞒ Ꞓ O S D C P T J K Pg N
	Fossil of Dickinsonia costata
	Fossil of Spriggina
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	†Proarticulata; Fedonkin, 1985
Classes and families
	†Vendiamorpha Fedonkin, 1985 †Vendiidae Ivantsov; ; †Dipleurozoa Harrington & Moore, 1955 †Dickinsoniidae ; Harrington & Moore, 1955; ; †Cephalozoa Ivantsov. 2004 †Sprigginidae Glaessner, 1958; †Yorgiidae Ivantsov, 2001; ; For more taxa, see text

Proarticulata is a proposed phylum of extinct, near-bilaterally symmetrical animals known from fossils found in the Ediacaran (Vendian) marine deposits, and dates to approximately 567 to 550 million years ago.^[1]^[2] The name comes from the Greek προ (pro-) = "before" and Articulata, i.e. prior to animals with true segmentation such as annelids and arthropods. This phylum was established by Mikhail A. Fedonkin in 1985 for such animals as Dickinsonia, Vendia, Cephalonega, Praecambridium^[3] and currently many other Proarticulata are described (see list).^[4]^[5]

Due to their simplistic morphology, their affinities and mode of life are subject to debate. They are almost universally considered to be metazoans, and due to possessing a clear central axis have been suggested to be stem-bilaterians. In the traditional interpretation, the Proarticulatan body is divided into transverse articulation (division) into isomers as distinct from the transverse articulation segments in annelids and arthropods, as their individual isomers occupy only half the width of their bodies, and are organized in an alternating pattern along the longitudinal axis of their bodies.^[5] In other words, one side is not the direct mirror image of its opposite (chirality). Opposite isomers of left and right side are located with displacement of half of their width. This phenomenon is described as the symmetry of gliding reflection.^[6]^[7] Some recent research suggests that some proarticulatans like Dickinsonia have genuine segments, and the isomerism is superficial and due to taphonomic distortion.^[8] However, other researchers dispute this.^[9]^[10] Displacement of left-right axis is known in bilaterians, notably lancelets.^[11]^[12]

Morphology

Vendiamorpha

The body is completely segmented, with all isomers curved towards the posterior, and the first isomer is normally much larger than the rest. The first two isomers at the anterior dorsal end are partly fused. (e.g., Vendia, Paravendia and Karakhtia).^[6]^[13]^[14]^[15]

Cephalozoa

These proarticulatans are incompletely segmented, as the anterior zone is free of isomers, often making a "hairband" like appearance (example cephalozoans include Yorgia, Praecambridium, Andiva, Archaeaspinus, Ivovicia, Podolimirus, Tamga, Spriggina, Marywadea and Cyanorus).^[6]^[13]^[15]^[16] Some cephalozoans from the family Yorgiidae demonstrate pronounced asymmetry of the left and right parts of the body. For instance, Yorgia’s initial right isomer is the only one which spreads far towards the left side of the body. Archaeaspinus has an unpaired anterior lobe confined by the furrow to the left side only.^[6]^[7]^[15]

In Cephalonega stepanovi and Tamga hamulifera the zone containing the isomers is encircled by a peripheral, undivided zone.^[16] The Cephalonega's isomers are connected to each other, forming a body resembling a rubber raft; the Tamga's isomers are separated from each other, and do not touch.

In Lossinia, the center undivided region has no visible isomers, instead having the lobe-like isomers emanate from the periphery of the undivided region as "transverse articulations."^[16]

Dipleurozoa

The dipleurozoan body is subradial, divided by isomers entirely (e.g., Dickinsonia and Phyllozoon). Dickinsonia juveniles show undivided anterior areas but these regions were reduced in the course of ontogeny, and in the adult stages Dickinsonia-like proarticulates changed so radically that they became almost indistinguishable from isomers.^[13]^[16]^[18]

List of Proarticulata

Andiva ivantsovi

Dickinsonia costata

Spriggina floundersi

Ovatoscutum concentricum

Yorgia waggoneri

Tamga hamulifera

Body fossils

Armillifera Fedonkin, 1980^[19]

A. parva Fedonkin, 1980

Andiva Fedonkin, 2002^[20]

A. ivantsovi Fedonkin, 2002

Archaeaspinus Ivantsov, 2007^[6] (=Archaeaspis Ivantsov, 2001)^[16]

A. fedonkini Ivantsov, 2001

Cephalonega Ivantsov et al., 2019^[21]

C. stepanovi (Fedonkin, 1976)^[16]^[22]

Chondroplon Wade, 1971 (possible =Dickinsonia)

C. bilobatum Wade, 1971

Cyanorus Ivantsov, 2004^[13]

C. singularis Ivantsov, 2004

Dickinsonia Sprigg, 1947

D. costata Sprigg, 1947

D. menneri Keller 1976^[16] (=Vendomia menneri Keller 1976^[22])

D. tenuis Glaessner & Wade, 1966

Ivovicia Ivantsov, 2007^[16]

I. rugulosa Ivantsov, 2007

Karakhtia Ivantsov, 2004

K. nessovi Ivantsov, 2004

Lossinia Ivantsov, 2007^[16]

L. lissetskii Ivantsov, 2007

Marywadea Glaessner, 1976

M. ovata Glaessner & Wade, 1966

Ovatoscutum Glaessner & Wade, 1966

O. concentricum Glaessner & Wade, 1966

Paravendia Ivantsov, 2004^[6]^[13]

P. janae Ivantsov, 2001 (=Vendia janae Ivantsov, 2001)

Podolimirus Fedonkin, 1983 (=Valdainia Fedonkin, 1983)^[23]

P. mirus Fedonkin, 1983 (Valdainia plumosa Fedonkin, 1983)

Praecambridium Glaessner & Wade, 1966

P. siggilum Glaessner & Wade, 1966

Spriggina Glaessner, 1958

S. floundersi Glaessner, 1958

Tamga Ivantsov, 2007^[16]

T. hamulifera Ivantsov, 2007

Vendia Keller, 1969^[6]^[13]

V. sokolovi Keller, 1969

V. rachiata Ivantsov, 2004

? Windermeria Narbonne, 1994

W. aitkeni Narbonne, 1994^[24]

Yorgia Ivantsov, 1999^[7]

Y. waggoneri Ivantsov, 1999

Trace fossils

Epibaion Ivantsov, 2002^[18]^[25]

E. axiferus Ivantsov, 2002.

E. waggoneris Ivantsov, 2011. This is a trace of Yorgia waggoneri

E. costatus Ivantsov, 2011. This is a trace of Dickinsonia costata

Phyllozoon Jenkins & Gehling, 1978

P. hanseni Jenkins & Gehling, 1978

References

^ Maslov AV, Podkovyrov VN, Grazhdankin DV, Kolesnikov AV (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 AV, Liu AG, Danelian T, Grazhdankin DV (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.
^ Fedonkin MA (1985). "Systematic Description of Vendian Metazoa". In Sokolov BS, Iwanowski AB (eds.). Vendian System: Historical–Geological and Paleontological Foundation. Vol. 1: Paleontology. Moscow: Nauka. pp. 70–106.
^ Fedonkin MA (31 March 2003). "The origin of the Metazoa in the light of the Proterozoic fossil record" (PDF). Paleontological Research. 7 (1): 9–41. doi:10.2517/prpsj.7.9. S2CID 55178329.
^ ^a ^b Ivantsov AY, Fedonkin MA, Nagovitsyn AL, Zakrevskaya ZA (2019). "Cephalonega, a new generic name, and the system of Vendian Proarticulata". Paleontological Journal. 53 (5): 447–454. doi:10.1134/S0031030119050046. S2CID 203853224.
^ ^a ^b ^c ^d ^e ^f ^g Ivantsov, A.Yu. (2001). "Vendia and other precambrian "Arthropods"". Paleontological Journal. 35 (4): 335–343.
^ ^a ^b ^c Ivantsov, A.Yu. (1999). "A new Dickinsoniid from the upper Vendian of the White Sea Winter Coast (Russia, Arkhangelsk region)". Paleontological Journal. 33 (3): 233–241.
^ Dunn FS, Liu AG, Donoghue PC (May 2018). "Ediacaran developmental biology". Biological Reviews of the Cambridge Philosophical Society. 93 (2): 914–932. doi:10.1111/brv.12379. PMC 5947158. PMID 29105292.
^ Ivantsov AY, Zakrevskaya MA, Nagovitsyn AL (June 2019). "Morphology of integuments of the Precambrian animals, Proarticulata". Invertebrate Zoology. 16 (1): 19–26. doi:10.15298/invertzool.16.1.03.
^ Ivantsov Y, Fedonkin MA, Nagovitsyn AL, Zakrevskaya MA (September 2019). "Cephalonega, A New Generic Name, and the System of Vendian Proarticulata". Paleontological Journal. 53 (5): 447–454. doi:10.1134/s0031030119050046. S2CID 203853224.
^ Blum M, Feistel K, Thumberger T, Schweickert A (April 2014). "The evolution and conservation of left-right patterning mechanisms". Development. 141 (8): 1603–13. doi:10.1242/dev.100560. PMID 24715452.
^ Soukup V (2017). "Left-right asymmetry specification in amphioxus: review and prospects". The International Journal of Developmental Biology. 61 (10–11–12): 611–620. doi:10.1387/ijdb.170251vs. PMID 29319110.
^ ^a ^b ^c ^d ^e ^f Ivantsov, A.Yu. (2004). "New Proarticulata from the Vendian of the Arkhangel'sk Region" (PDF). Paleontological Journal. 38 (3): 247–253. Archived from the original (PDF) on 2007-09-27.
^ Ivantsov, A.Yu.; Malakhovskaya, Y.E.; Serezhnikova, E.A. (2004). "Some problematic fossils from the Vendian of the south-eastern White Sea region" (PDF). Paleontological Journal. 38 (1): 1–9. Archived from the original (PDF) on 2007-07-04.
^ ^a ^b ^c Ivantsov, A.Yu. (2004). Vendian animals in the phylum Proarticulata (PDF). The Rise and Fall of the Vendian Biota. Prato, Italy. p. 52. IGSP Project 493.
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j Ivantsov, A.Yu. (April 2007). "Small Vendian transversely articulated fossils". Paleontological Journal. 41 (2): 113–122. doi:10.1134/S0031030107020013. S2CID 86636748.
^ illustration (c) Stanton F. Fink
^ ^a ^b Ivantsov, A.Yu.; Malakhovskaya, Y.E. (2002). "Giant traces of Vendian animals" (PDF). Doklady Earth Sciences. 385 (6): 618–622. Archived from the original (PDF) on 2007-07-04.
^ Ivantsov AY (December 2010). "Paleontological evidence for the supposed precambrian occurrence of mollusks". Paleontological Journal. 40 (12): 1552–1559. doi:10.1134/S0031030110120105. S2CID 86523806.
^ Fedonkin MA (2002). "Andiva ivantsovi gen. et sp. n. and related carapace‐bearing Ediacaran fossils from the Vendian of the Winter Coast, White Sea, Russia". Italian Journal of Zoology. 69 (2): 175–181. doi:10.1080/11250000209356456. S2CID 85352552.
^ Ivantsov AY, Fedonkin MA, Nagovitsyn AL, Zakrevskaya MA (2019). "Cephalonega, a new generic name, and the system of Vendian Proarticulata". Paleontological Journal. 53 (5): 447–454. doi:10.1134/S0031030119050046. S2CID 203853224.
^ ^a ^b Keller BM, Fedonkin MA (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 2007-09-27.
^ Dzik, J.; Martyshyn, A. (2015). "Taphonomy of the Ediacaran Podolimirus and associated dipleurozoans from the Vendian of Ukraine". Precambrian Research. 269: 139–146. doi:10.1016/j.precamres.2015.08.015.
^ Narbonne GM (May 1994). "New Ediacaran fossils from the Mackenzie Mountains, Northwestern Canada". Journal of Paleontology. 63 (3): 411–416. doi:10.1017/S0022336000025816. JSTOR 1306192. S2CID 133215767.
^ Ivantsov AY (May 2011). "Feeding traces of Proarticulata — the Vendian metazoa". Paleontological Journal. 45 (3): 237–248. doi:10.1134/S0031030111030063. S2CID 128741869.

External links

Database of Ediacaran Biota Advent of Complex Life

[Maslov_2018-1] Maslov AV, Podkovyrov VN, Grazhdankin DV, Kolesnikov AV (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_2018-2] Kolesnikov AV, Liu AG, Danelian T, Grazhdankin DV (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.

[Fedonkin_1985-3] Fedonkin MA (1985). "Systematic Description of Vendian Metazoa". In Sokolov BS, Iwanowski AB (eds.). Vendian System: Historical–Geological and Paleontological Foundation. Vol. 1: Paleontology. Moscow: Nauka. pp. 70–106.

[Fedonkin2003-4] Fedonkin MA (31 March 2003). "The origin of the Metazoa in the light of the Proterozoic fossil record" (PDF). Paleontological Research. 7 (1): 9–41. doi:10.2517/prpsj.7.9. S2CID 55178329.

[Ivantsov_2019-5] Ivantsov AY, Fedonkin MA, Nagovitsyn AL, Zakrevskaya ZA (2019). "Cephalonega, a new generic name, and the system of Vendian Proarticulata". Paleontological Journal. 53 (5): 447–454. doi:10.1134/S0031030119050046. S2CID 203853224.

[Ivantsov2001-6] ^ ^a ^b ^c ^d ^e ^f ^g Ivantsov, A.Yu. (2001). "Vendia and other precambrian "Arthropods"". Paleontological Journal. 35 (4): 335–343.

[Ivantsov1999-7] Ivantsov, A.Yu. (1999). "A new Dickinsoniid from the upper Vendian of the White Sea Winter Coast (Russia, Arkhangelsk region)". Paleontological Journal. 33 (3): 233–241.

[8] Dunn FS, Liu AG, Donoghue PC (May 2018). "Ediacaran developmental biology". Biological Reviews of the Cambridge Philosophical Society. 93 (2): 914–932. doi:10.1111/brv.12379. PMC 5947158. PMID 29105292.

[9] Ivantsov AY, Zakrevskaya MA, Nagovitsyn AL (June 2019). "Morphology of integuments of the Precambrian animals, Proarticulata". Invertebrate Zoology. 16 (1): 19–26. doi:10.15298/invertzool.16.1.03.

[10] Ivantsov Y, Fedonkin MA, Nagovitsyn AL, Zakrevskaya MA (September 2019). "Cephalonega, A New Generic Name, and the System of Vendian Proarticulata". Paleontological Journal. 53 (5): 447–454. doi:10.1134/s0031030119050046. S2CID 203853224.

[11] Blum M, Feistel K, Thumberger T, Schweickert A (April 2014). "The evolution and conservation of left-right patterning mechanisms". Development. 141 (8): 1603–13. doi:10.1242/dev.100560. PMID 24715452.

[12] Soukup V (2017). "Left-right asymmetry specification in amphioxus: review and prospects". The International Journal of Developmental Biology. 61 (10–11–12): 611–620. doi:10.1387/ijdb.170251vs. PMID 29319110.

[Ivantsov2004-13] ^ ^a ^b ^c ^d ^e ^f Ivantsov, A.Yu. (2004). "New Proarticulata from the Vendian of the Arkhangel'sk Region" (PDF). Paleontological Journal. 38 (3): 247–253. Archived from the original (PDF) on 2007-09-27.

[Ivantsov2004et_al-14] Ivantsov, A.Yu.; Malakhovskaya, Y.E.; Serezhnikova, E.A. (2004). "Some problematic fossils from the Vendian of the south-eastern White Sea region" (PDF). Paleontological Journal. 38 (1): 1–9. Archived from the original (PDF) on 2007-07-04.

[Ivantsov2004Prato-15] Ivantsov, A.Yu. (2004). Vendian animals in the phylum Proarticulata (PDF). The Rise and Fall of the Vendian Biota. Prato, Italy. p. 52. IGSP Project 493.

[Ivantsov2007-16] ^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j Ivantsov, A.Yu. (April 2007). "Small Vendian transversely articulated fossils". Paleontological Journal. 41 (2): 113–122. doi:10.1134/S0031030107020013. S2CID 86636748.

[17] ustration (c) Stanton F. Fink

[Ivantsov2002-18] Ivantsov, A.Yu.; Malakhovskaya, Y.E. (2002). "Giant traces of Vendian animals" (PDF). Doklady Earth Sciences. 385 (6): 618–622. Archived from the original (PDF) on 2007-07-04.

[Ivantsov2010-19] Ivantsov AY (December 2010). "Paleontological evidence for the supposed precambrian occurrence of mollusks". Paleontological Journal. 40 (12): 1552–1559. doi:10.1134/S0031030110120105. S2CID 86523806.

[Fedonkin2002-20] Fedonkin MA (2002). "Andiva ivantsovi gen. et sp. n. and related carapace‐bearing Ediacaran fossils from the Vendian of the Winter Coast, White Sea, Russia". Italian Journal of Zoology. 69 (2): 175–181. doi:10.1080/11250000209356456. S2CID 85352552.

[21] Ivantsov AY, Fedonkin MA, Nagovitsyn AL, Zakrevskaya MA (2019). "Cephalonega, a new generic name, and the system of Vendian Proarticulata". Paleontological Journal. 53 (5): 447–454. doi:10.1134/S0031030119050046. S2CID 203853224.

[Keller_Fedonkin_1976-22] Keller BM, Fedonkin MA (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 2007-09-27.

[Dzik2015-23] Dzik, J.; Martyshyn, A. (2015). "Taphonomy of the Ediacaran Podolimirus and associated dipleurozoans from the Vendian of Ukraine". Precambrian Research. 269: 139–146. doi:10.1016/j.precamres.2015.08.015.

[Narbonne1994-24] Narbonne GM (May 1994). "New Ediacaran fossils from the Mackenzie Mountains, Northwestern Canada". Journal of Paleontology. 63 (3): 411–416. doi:10.1017/S0022336000025816. JSTOR 1306192. S2CID 133215767.

[Ivantsov2011-25] Ivantsov AY (May 2011). "Feeding traces of Proarticulata — the Vendian metazoa". Paleontological Journal. 45 (3): 237–248. doi:10.1134/S0031030111030063. S2CID 128741869.

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PROFILPELAJAR.COM

Proarticulata

Morphology

Vendiamorpha

Cephalozoa

Dipleurozoa

List of Proarticulata

Body fossils

Trace fossils

See also

References

External links

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