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Ctenocystoidea is an extinct clade of
echinoderm An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the s ...
s, which lived during the Cambrian and
Ordovician The Ordovician ( ) is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start of the Silurian Period Mya. T ...
periods. Unlike other echinoderms, ctenocystoids had bilateral symmetry, or were only very slightly asymmetrical. They are believed to be one of the earliest-diverging branches of echinoderms, with their bilateral symmetry a trait shared with other deuterostomes. Ctenocystoids were once classified in the taxon
Homalozoa Homalozoa is an obsolete extinct subphylum of Paleozoic era echinoderms, prehistoric marine invertebrates. They are also referred to as carpoids. Description The Homalozoa lacked the typical pentamer body form of other echinoderms, but all were ...
, also known as Carpoidea, alongside cinctans,
solutes In chemistry, a solution is a special type of homogeneous mixture composed of two or more substances. In such a mixture, a solute is a substance dissolved in another substance, known as a solvent. If the attractive forces between the solvent ...
, and stylophorans. Homalozoa is now recognized as a polyphyletic group of echinoderms without radial symmetry. Ctenocystoids were geographically widespread during the
Middle Cambrian Middle or The Middle may refer to: * Centre (geometry), the point equally distant from the outer limits. Places * Middle (sheading), a subdivision of the Isle of Man * Middle Bay (disambiguation) * Middle Brook (disambiguation) * Middle Creek ...
, with one species surviving into the
Late Ordovician The Ordovician ( ) is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start of the Silurian Period Mya. ...
.


Description

Like other echinoderms, ctenocystoids had a skeleton made of stereom plates. Ctenocystoids had near bilateral symmetry, with some species exhibiting slightly asymmetric plate shapes. Like cinctans and ''
Ctenoimbricata ''Ctenoimbricata'' is an extinct genus of bilaterally symmetrical echinoderm, which lived during the early Middle Cambrian period of what is now Spain. It contains one species, ''Ctenoimbricata spinosa''. It may be the most basal known echinoder ...
'', most ctenocystoids had large marginal plates surrounding the sides of their body, but unlike cinctans and ''Ctenoimbricata'', which had only one row of marginal plates, most ctenocystoids had two rows of marginal plates. ''Courtessolea'' had only one row of marginal plates, like cinctans and ''Ctenoimbricata'', whereas ''Conollia'' lost the marginal plates entirely. The anus of ctenocystoids was surrounded by a pyramidal
periproct The periproct is the final body segment in annelid worms. The anus is located on this segment. The term also refers to the small region surrounding the anus of the sea urchin. See also *Prostomium *Earthworm *Sea urchin Sea urchins () are s ...
as in other echinoderms. It was located at the posterior end, defining a clear anterior-posterior body axis unlike other echinoderms. All ctenocystoids had a ctenoid apparatus, a comb-like arrangement of movable plates at the anterior end of the animal. Beneath the ctenoid apparatus was a mouth, bordered on each side by a food groove.


Classification


Relationships

Ctenocystoids are likely among the most basal stem-group echinoderms. They have also been interpreted as aberrant
blastozoa Blastozoa is a subphylum of extinct animals belonging to Phylum Echinodermata. This subphylum is characterized by the presence of specialized respiratory structures and brachiole plates used for feeding. This subphylum ranged from the Cambrian to ...
ns and as stem-group
hemichordates Hemichordata is a phylum which consists of triploblastic, enterocoelomate, and bilaterally symmetrical marine deuterostome animals, generally considered the sister group of the echinoderms. They appear in the Lower or Middle Cambrian and includ ...
. The presence of stereom plates indicates that they most likely belong to the echinoderm total group, rendering a hemichordate affinity unlikely. ''Courtessolea'' was probably the most basal ctenocystoid, given its anatomical similarities to ''Ctenoimbricata'' and cinctans. ''Conollia'' and ''Jugoszovia'' may be closely related to each other, as both have a reduced marginal frame compared to other ctenocystoids.


Genera

The following genera of ctenocystoids have been named:


Distribution

Ctenocystoids were widespread during the Middle Cambrian, and have been found in the United States, the United Kingdom, Australia, France, the Czech Republic, Poland, Spain, and Morocco. The earliest ctenocystoids date to the beginning of Stage 5 of the Cambrian, now known as the
Wuliuan The Wuliuan stage is the fifth stage of the Cambrian, and the first stage of the Miaolingian Series of the Cambrian. It was formally defined by the ICS in 2018. Its base is defined by the first appearance of the trilobite species '' Oryctoceph ...
age, or possibly slightly earlier, in late Cambrian Stage 4. Most species date to the Wuliuan and
Drumian The Drumian is a stage of the Miaolingian Series of the Cambrian. It succeeds the Wuliuan and precedes the Guzhangian. The base is defined as the first appearance of the trilobite '' Ptychagnostus atavus'' around million years ago. The top i ...
ages of the Cambrian. The geologically youngest ctenocystoid, the only one known from the
Ordovician The Ordovician ( ) is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start of the Silurian Period Mya. T ...
, is ''Conollia'', from the
Sandbian The Sandbian is the first stage of the Upper Ordovician. It follows the Darriwilian and is succeeded by the Katian. Its lower boundary is defined as the first appearance datum of the graptolite species '' Nemagraptus gracilis'' around million yea ...
of the United Kingdom.


History

The class Ctenocystoidea was named in 1969 by Richard A. Robison and James Sprinkle. It originally contained one species, ''Ctenocystis utahensis''. The name comes from the Greek words '' ktenos'', meaning "comb", and '' kystis'', meaning "sac". It was originally assigned to the echinoderm subphylum
Homalozoa Homalozoa is an obsolete extinct subphylum of Paleozoic era echinoderms, prehistoric marine invertebrates. They are also referred to as carpoids. Description The Homalozoa lacked the typical pentamer body form of other echinoderms, but all were ...
.


References

{{Taxonbar, from=Q5792856 Paleozoic echinoderms Prehistoric deuterostome classes Homalozoa