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Proarticulata is a of extinct, near-bilaterally symmetrical known from fossils found in the (Vendian) marine deposits, and dates to approximately . The name comes from the Greek προ () = "before" and Articulata, i.e. prior to animals with true segmentation such as and . This phylum was established by Mikhail A. Fedonkin in 1985 for such animals as , , , and currently many other Proarticulata are described (see list).

Due to their simplistic morphology, their affinities and mode of life are subject to debate. They are almost universally considered to be , and due to possessing a clear central axis have been suggested to be stem-. 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. In other words, one side is not the direct mirror image of its opposite ( ). Opposite isomers of left and right side are located with displacement of half of their width. This phenomenon is described as the symmetry of . Some recent research suggests that some proarticulatans like Dickinsonia have genuine segments, and the isomerism is superficial and due to . However, other researchers dispute this. Displacement of left-right axis is known in bilaterians, notably .


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., , and ).


Cephalozoa
These proarticulatans are incompletely segmented, as the anterior zone is free of isomers, often making a "hairband" like appearance (example cephalozoans include , , , , , , Tamga, , and ).

Some cephalozoans from the family demonstrate pronounced asymmetry of the left and right parts of the body. For instance, Yorgia 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.

In stepanovi and the zone containing the isomers is encircled by a peripheral, undivided zone. The isomers are connected to each other, forming a body resembling a rubber raft; the isomers are separated from each other, and do not touch.

In , 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."


Dipleurozoa
The dipleurozoan body is subradial, divided by isomers entirely (e.g., and ). 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.


List of Proarticulates

Body fossils
A. parva Fedonkin, 1980
A. ivantsovi Fedonkin, 2002
A. fedonkini Ivantsov, 2001
C. stepanovi (Fedonkin, 1976)
C. bilobatum Wade, 1971
C. singularis Ivantsov, 2004
D. costata Sprigg, 1947
D. menneri Keller 1976 (= Vendomia menneri Keller 1976)
D. tenuis Glaessner & Wade, 1966
I. rugulosa Ivantsov, 2007
K. nessovi Ivantsov, 2004
L. lissetskii Ivantsov, 2007
M. ovata Glaessner & Wade, 1966
O. concentricum Glaessner & Wade, 1966
P. janae Ivantsov, 2001 (= Vendia janae Ivantsov, 2001)
  • Fedonkin, 1983 (= Valdainia Fedonkin, 1983)
P. mirus Fedonkin, 1983 ( Valdainia plumosa Fedonkin, 1983)
P. siggilum Glaessner & Wade, 1966
S. floundersi Glaessner, 1958
  • Tamga Ivantsov, 2007
T. hamulifera Ivantsov, 2007
V. sokolovi Keller, 1969
V. rachiata Ivantsov, 2004
W. aitkeni Narbonne, 1994
Y. waggoneri Ivantsov, 1999


Trace fossils
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
P. hanseni Jenkins & Gehling, 1978


See also
  • Articulata
  • List of Ediacaran genera


External links

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