Isomer (Proarticulata)
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Isomer (Greek ''isos'' = "equal", ''méros'' = "part") is an element of transverse body articulation of the bilateral fossil animals of the
Phylum In biology, a phylum (; plural: phyla) is a level of classification or taxonomic rank below kingdom and above class. Traditionally, in botany the term division has been used instead of phylum, although the International Code of Nomenclature f ...
Proarticulata Proarticulata is a proposed phylum of extinct, bilaterally symmetrical animals known from fossils found in the Ediacaran (Vendian) marine deposits, and dates to approximately . The name comes from the Greek () = "before" and Articulata, i.e. pr ...
from the
Ediacaran The Ediacaran Period ( ) is a geological period that spans 96 million years from the end of the Cryogenian Period 635 million years ago (Mya), to the beginning of the Cambrian Period 538.8 Mya. It marks the end of the Proterozoic Eon, and th ...
(Vendian) period. This term has been proposed by Andrey Yu. Ivantsov, a Russian paleontologist from the Laboratory of the Precambrian organisms,
Paleontological Institute, Russian Academy of Sciences The Paleontological Institute, Russian Academy of Sciences (PIN; russian: Палеонтологический институт РАН) in Moscow is among the world's largest paleontological institutes. An affiliate of the Russian Academy of Scienc ...
.


Morphology

Proarticulatan isomers are distinct from the segments of the Annelida and
Panarthropoda Panarthropoda is a proposed animal clade containing the extant phyla Arthropoda, Tardigrada (water bears) and Onychophora (velvet worms). Panarthropods also include extinct marine legged worms known as lobopodians ("Lobopodia"), a paraphylet ...
, as each of these elements occupies only half of width of a body and are organized in an alternating pattern relatively to the axis of the body. In other words, although proarticulatans are bilaterally symmetrical, 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 its width. This phenomenon is described as the symmetry of gliding reflection.M. A. Fedonkin (1985). "Systematic Description of Vendian Metazoa". In Sokolov, B. S. and Iwanowski, A. B., eds., "Vendian System: Historical–Geological and Paleontological Foundation, Vol. 1: Paleontology". Moscow: Nauka, pp. 70–106. However, this type of symmetry is not unique to Proarticulata, a modern
lancelet The lancelets ( or ), also known as amphioxi (singular: amphioxus ), consist of some 30 to 35 species of "fish-like" benthic filter feeding chordates in the order Amphioxiformes. They are the modern representatives of the subphylum Cephalochorda ...
s have analogous asymmetric arrangement of
myomere Myomeres are blocks of skeletal muscle tissue arranged in sequence, commonly found in aquatic chordates. Myomeres are separated from adjacent myomeres by connective fascia (myosepta) and most easily seen in larval fishes or in the olm. Myomere ...
s and
somites The somites (outdated term: primitive segments) are a set of bilaterally paired blocks of paraxial mesoderm that form in the embryonic stage of somitogenesis, along the head-to-tail axis in segmented animals. In vertebrates, somites subdivide i ...
of
larvae A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
. The first element in the row is a right isomer on the anterior end of the dorsal side. All successive isomers are similar to it, but gradually decrease in size and in angle of inclination from the anterior part (head) of the body to the posterior end. New isomers were added at the growth point at the posterior end of the proarticulate body. With age, the addition of new isomers slowed down and probably stopped, the growth point shifted from the posterior end, and the relative length of the posterior isomers increased, sometimes significantly. The overall number of body isomers amounts from several pairs (''
Vendia ''Vendia'' is a genus of oval-shaped, Ediacaran fossils ranging from 4.5 to 12.5 mm long. The body is completely segmented into isomers, which are arranged alternately in two rows longitudinal to the axis of the body. The larger isomers cov ...
'', '' Onega'') to several hundreds (''
Dickinsonia ''Dickinsonia'' is an extinct genus of basal animal that lived during the late Ediacaran period in what is now Australia, China, Russia and Ukraine. The individual ''Dickinsonia'' typically resembles a bilaterally symmetrical ribbed oval. Its a ...
'').


Variation of Articulation Across Classes


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 together to form a headshield-like structure. (e.g., ''
Vendia ''Vendia'' is a genus of oval-shaped, Ediacaran fossils ranging from 4.5 to 12.5 mm long. The body is completely segmented into isomers, which are arranged alternately in two rows longitudinal to the axis of the body. The larger isomers cov ...
'', ''
Paravendia ''Paravendia'' is an extinct genus of proarticulate vendiamorph that lived in the Ediacaran period, about 553 million years ago. It shares the Vendiidae family with ''Vendia'' and ''Karakhtia''. It is a monotypic genus, with the species ''Pa ...
'', and ''
Karakhtia ''Karakhtia nessovi'' is a species of Proarticulate from the Ediacaran period, around 555 Million Years Ago. K. nessovi is the only species in the genus ''Karakhtia''. The genus Haootia has been compared minorly to ''Karakhtia'' in the way t ...
'').Ivantsov, A.Y. (2004
"Vendian Animals in the Phylum Proarticulata"
. The Rise and Fall of the Vendian Biota. IGSP Project 493. Abstracts. Prato, Italy, p. 52.


Cephalozoa (preliminary name)

These proarticulatans demonstrate incomplete segmentation, as the anterior zone is free of isomers, often making a "hairband" like appearance. (example cephalozoans include ''
Yorgia ''Yorgia waggoneri'' is a discoid Ediacaran organism. It has a low, segmented body consisting of a short wide "head", no appendages, and a long body region, reaching a maximum length of . It is classified within the extinct animal phylum Proarti ...
'', ''
Praecambridium ''Praecambridium sigillum'' is an extinct organism that superficially resembles a segmented trilobite-like arthropod. It was originally described as being a trilobite-like arthropod, though the majority of experts now place it within the Proartic ...
'', ''
Andiva ''Andiva ivantsovi'' is a Vendian fossil, identified to be a bilaterian triploblastic animal in the Ediacaran phylum Proarticulata, known from the Winter Coast, White Sea, Russia. It was first discovered in 1977, and described as a new species i ...
'', ''
Archaeaspinus ''Archaeaspinus fedonkini'' is an extinct proarticulatan organism from the Late Precambrian (Ediacaran) period. Background ''Archaeaspinus'' was discovered in Zimnii Bereg, the Winter Coast of the White Sea in Russia, by A. Yu. Ivantsov in 2001 ...
'', '' Ivovicia'', ''
Spriggina ''Spriggina'' is a genus of early bilaterian animals whose relationship to living animals is unclear. Fossils of ''Spriggina'' are known from the late Ediacaran period in what is now South Australia. ''Spriggina floundersi'' is the official fo ...
'', '' Marywadea'' and '' Cyanorus''. Some cephalozoans from the
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Yorgiidae Yorgiidae is an extinct family of cephalozoans, which lived 635 million years ago. They were filter fed. Description Like most proarticulates, they present semi-bilateral symmetry. They had a discoid appearance, with the body segmented by isome ...
demonstrate pronounced asymmetry of 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.


Dipleurozoa

The dipleurozoan body is subradial divided by isomers entirely (e.g., ''
Dickinsonia ''Dickinsonia'' is an extinct genus of basal animal that lived during the late Ediacaran period in what is now Australia, China, Russia and Ukraine. The individual ''Dickinsonia'' typically resembles a bilaterally symmetrical ribbed oval. Its a ...
'' and ''
Phyllozoon ''Phyllozoon''(lit. "Leaf animal" in greek) is an Ediacaran imprint that resembles a proarticulatan and has been interpreted as a feeding trace. It usually occurs in long chains of imprints formed, presumably as the organism that made it moved. ...
''). Juveniles ''Dickinsonia'' show an undivided anterior area but this region was reduced in the course of ontogeny, and in adult ''Dickinsonia''-like proarticulates changed so strongly that became almost indistinguishable from isomers.


Proarticulata ''Incertae sedis''

In ''
Onega stepanovi ''Cephalonega stepanovi'' is a fossil organism from Ediacaran deposits of the Arkhangelsk Region, Russia. It was described by Mikhail A. Fedonkin in 1976 Name Its original genus name ''Onega'' comes from the Onega Peninsula of the White Sea, ...
'' and '' Tamga hamulifera'' all isomers are encircled by a peripheral undivided zone. In the former, the isomers remain in contact with each other, while in the latter, the isomers do not touch.


See also

*
Sea pen Sea pens are colonial marine cnidarians belonging to the order Pennatulacea. There are 14  families within the order; 35 extant genera, and it is estimated that of 450 described species, around 200 are valid. Sea pens have a co ...
- extant order with some members exhibiting glide reflection symmetry


References

{{reflist Proarticulata Animal anatomy