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Ambulacraria

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Echinoderms are the sister group of the Hemichordata, with which they form the crown group Ambulacraria.[1] Two taxa of uncertain placement, Vetulocystida and Yanjiahella, have each been proposed as either stem-group echinoderms[2][3] or stem-group ambulacrarians.[4][5] Vetulocystids have also been proposed as stem-group chordates,[6] while Yanjiahella has also been proposed to be a stem-group hemichordate.[5]

The Ambulacrarian context of the echinoderms is shown below, simplified from Li et al. 2023,[7] with the possible ambulacrarian placements of the uncertian taxa shown with dashed lines and question marks:

Xenambulacraria

? Xenacoelomorpha

Ambulacraria
Vetulocystida †

   

Cambroernida †

   

Yanjiahella †

   

Ambulacraria
Echinodermata
? Vetulocystida †

   

Yanjiahella †

   

Echinodermata

   

Hemichordata
Yanjiahella †

   

Hemichordata

   

Chordata
? Vetulocystida †

   

Chordata

   

Echinoderms

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Material for Later

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Asterozoa

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Asterozoa

Cantabrigiaster

Villebrunaster

Chinianaster

Thoralaster

Asterozoa

Ophioxenikos

Archegonaster

Phragmactis

Rhopalocoma

Maydenia

Eriniceaster

Pradesura

Eophiura

Palaeura

Protaster

Encrinaster

Ophiocantabria   

Stenaster

Urasterella

Cnemidactis

Platanaster

Hudsonaster

Petraster

(crown group)
(total group)
Somasteroidea
Stenuroidea
Ophiuroidea
Asteroidea

Crinozoa: External phylogeny

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Echinodermata
Echinodermata

? Arkarua

Helicoplacoidea

Edrioasteroidea

Blastozoa
(including Ctenocystoidea, Cincta, & Soluta)

Pelmatozoa and Eleutherozoa

F. A. Bather produced the earliest widely used classification of both fossil and extant echinoderms in 1900.[8] His primary division was between Pelmatozoa and Eleutherozoa, both of which continue to be referenced in cladistic work despite being discarded by later taxonomies.[9][10]

Pelmatozoa was originally defined by Leuckart for stemmed groups, but by 1900 had come to be used for all immobile forms. Eleutherozoa contained all mobile forms, which Bather believed had each descended from Edrioasteroidean groups at different times.[11] In cladistic terms, this made Bather's Pelmatozoa paraphyletic with respect to a polyphyletic Eleutherozoa.

Ironically, cladistic approaches have consistently recovered Eleutherozoa as a monophyletic clade, a view now considered "uncontroversial."[12] There is no such consensus regarding Pelmatozoa. Recent work restricts Pelmatozoa to its original scope of stemmed groups, but has differed as to whether it is monophyletic,[13][14], paraphyletic,[15] ambiguous,[16] or simply useful as an informal term describing body plans.[17]

Homalozoa, Crinozoa, Echinozoa, and Asterozoa

In 1966 the Treatise on Invertebrate Paleontology, edited by Raymond Cecil Moore, rejected Bather's classification, replacing it with a new four-subphylum scheme[18] that had been previously proposed by H. B. Fell.[19] In addition to the obvious changes, the contents of Bather's Cystidea were distributed among the three Homalozoan classes and various Crinozoan classes, particularly Cystoidea and Eocrinoidea.[20]

Artist's reconstruction of Camptostroma

Of these divisions, Camptostromatoidea is the only one not addressed by Sprinkle's subsequent revision (discussed below), although he notes that its sole genus, Camptostroma, is "strange."[21] Subsequent research has placed Camptostroma as a basal Edrioasteroid.[22]


Adding Blastozoa as a fifth subphylum

Difference between crinoid arms, which are extensions of the main body, and blastoid brachioles, which are not, had been observed for some time. In 1973, James Sprinkle concluded that enough fossil evidence of brachiole-bearing echinoderms had been assembled to justify placing them in their own phylum, Blastozoa, restricting Crinozoa to stalked forms with arms.[23]

arms vs brachioles[24]

James Sprinkle which added a fifth subphylum to the Treatise taxonomy in 1973.[25] His later class-level taxonomy of the five subphyla was the most recent approach cited in an early cladistic re-assessment of the phylum.[26]


Sprinkle split Blastozoa, defined by the presence of brachioles, out of Crinozoa, which he redefined as stalked taxa with arms.[27] He also moved Edrioblastoidea to the Echinozoa[28], replaced Cystoidea by promoting its two orders, Rhombifera and Diploporita, to class level, and similarly replaced Stellaroidea by promoting two of its sublasses, Asteroidea and Ophiuroidea.[29] Finally, he demoted Lepidocystoidea, moving it under Eocrinoidea[30], and dropped Camptostromatoidea as a class without explicitly reassigning the "strange" genus Camptostroma.[31]

In addition to the obvious re-allocation of formerly Crinozoan classes to Blastozoa and (in the case of Edrioblastoidea) Echinozoa, Sprinkle made the following changes:

  • Demoted Lepidocystoidea and placed it under Eocrinoidea[32]
  • Demoted Camptostromatoidea without placing its "strange" sole genus, Camptostroma, into another class[33]
  • Adopted Christopher R. C. Paul's 1968 raising of Rhombifera and Diploporita to class level, replacing Cystoidea which had contained only those two taxa as orders[34]
  • Adopted the replacement of Stelleroidea with two of its three sub-classes, Asteroidea and Ophiuroidea, promoted to class level

The division of classes between Crinozoa and Blastozoa, and the relationship of the subphyla to each other, remains a substantial point of debate. Some workers have found Blastozoa to be paraphyletic with respect to Crinozoa,Cite error: A <ref> tag is missing the closing </ref> (see the help page). or polyphyletic,[35] as illustrated by cladograms in the next section.

Echinozoa and Asterozoa have been accepted as monophyletic sub-taxa of Eleutherozoa,[36][37] although a few workers continue to support the clade Cryptosyringida, uniting Ophiuroida and Echinozoa as the sister group of the Asteroidea.

Under cladistic approaches, Eleutherozoa and Asterozoa have been proven to be monophyletic as defined above, as have Crinoidea and a reduced Echinozoa limited to Echinoidea, Holothuroidea, and Ophiocistioidea.[38][39][40][41]

While each class within the Homolozoa is still thought to be monophyletic, the subphylum itself is either polyphyletic[42] or paraphyletic.[43] The contents, relationships, and monophyly of Blastozoa, Crinozoa (beyond Crinoidea), Pelmatozoa (often narrowed to include only stemmed classes), their constituent classes, and the classes excluded from the narrowed Echinozoa, remain contentious.[44][45][46][47][48][49][50][51][52]

probably not

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Bather also provided a diagram[53] that, together with his statement that "even if all Eleutherozoa descended from one class of Pelmatozoa, they did so at widely differing periods" and in a specific order,[54] is roughly equivalent to the following cladogram:

Pelmatozoa

Cystidea†

(Homalozoans, Eocrinoids, Paracrinoids, Cystoids)[55]

Blastoidea†

Crinoidea

some Edrioasteroids†

Edrioasteroidea

Holothuroidea

some Edrioasteroids†

Echinoidea

some Edrioasteroids†  

Stelleroidea

Eleutherozoa

This shows Pelmatozoa and Edrioasteroidea as paraphyletic with respect to a polyphyletic Eleutherozoa


Internal phylogeny
Echino‑

Crinoidea (feather stars)

Echinozoa
Holothuroidea

sea cucumbers
Echinoidea

sea urchins, etc
Asterozoa
Ophiuroidea

brittle & basket stars
Asteroidea

starfish
dermata

Fossil history

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Linnean Taxonomy

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Phylogeny

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David et. al 2000

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The following cladogram is based on David & Mooi (1999)[56] and David, Lefebvre, Mooi, and Parsley (2000):[57]

Notes:

  • Parablastoidea and Paracrinoidea are stated to be part of Blastozoa, but are not placed on the cladogram[58]
  • Edrioblastoidea is not mentioned, and while the sole genus Camptostroma of Camptostromatoidea is mentioned, it is not explicitly associated with a class
  • Helicocystis, Ctenoimbricata, and Yanjiahella had not yet been discovered (see next cladogram for their significance)
  • Pelmatozoa is shown to be paraphyletic[59]

Zamora & Smith 2024

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The following cladogram is based on Rahman & Zamora (2024),[60], using class names from the cladogram's caption,[61] plus additional subphylum names[62] from the article text. A dashed line and a question mark indicate uncertain placement:

Echinodermata
   

Ctenoimbricata

Ctenocystoidea

   

Cincta(Homostelea)

   
   

Soluta(Homoiostelea)

Stylophora† ?

Notes

  • Ophiocistioidea is not addressed by this paper, see the previous cladogram above for a typical placement
  • Non-eocrinoid blastozoans are mentioned but not placed on the cladogram
  • Arkarua is not considered an echinoderm and is intentionally omitted[63]

Bather 1900

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Pelmatozoa
Cystidea†

some Eocrinoids†

some Eocrinoids† + some Cornute Stylophorans†

Homostelea† (Cincta) + some Cornute Stylophorans†

Mitrate Stylophorans†

Homoiostelea† (Soluta)

some Diploporitans†

Cystoidea†

some Diploporitans†

Rhombifera†

Blastoidea

Crinoidea

Edrioasteroidea

some Edrioasteroids

Eleutherozoa

Holothuroidea

some Edrioasteroids

Echinoidea

some Edrioasteroids

Stelleroidea

Asteroidea

Ophiuroidea

Smith 1984

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Smith revived the name "Cystoidea" in place of Sprinkle's "Blastozoa", including the traditional Eocrinoidea, Rhombifera, Diploporita, Blastoidea, and Paracrinoidea within it.[64]

Existing Text

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The oldest candidate echinoderm fossil is Arkarua from the Precambrian of Australia. These fossils are disc-like, with radial ridges on the rim and a five-pointed central depression marked with radial lines. However, the fossils have no stereom or internal structure indicating a water vascular system, so they cannot be conclusively identified.[65]

The first universally accepted echinoderms appear in the Lower Cambrian period; asterozoans appeared in the Ordovician, while the crinoids were a dominant group in the Paleozoic. Echinoderms left behind an extensive fossil record.[66] It is hypothesised that the ancestor of all echinoderms was a simple, motile, bilaterally symmetrical animal with a mouth, gut and anus. This ancestral organism adopted an attached mode of life with suspension feeding, and developed radial symmetry. Even so, the larvae of all echinoderms are bilaterally symmetrical, and all develop radial symmetry at metamorphosis. Like their ancestor, the starfish and crinoids still attach themselves to the seabed while changing to their adult form.[67]

The first echinoderms were non-motile,[67] but evolved into animals able to move freely. These soon developed endoskeletal plates with stereom structure, and external ciliary grooves for feeding.[68] The Paleozoic echinoderms were globular, attached to the substrate and were orientated with their oral surfaces facing upwards. These early echinoderms had ambulacral grooves extending down the side of the body, fringed on either side by brachioles, like the pinnules of a modern crinoid. Eventually, except for the crinoids, all the classes of echinoderms reversed their orientation to become mouth-downward. Before this happened, the podia probably had a feeding function, as they do in the crinoids today. The locomotor function of the podia came later, when the re-orientation of the mouth brought the podia into contact with the substrate for the first time.[67]

Further information: Dibrachicystis

Alternative echinoderm trees

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Rahman & Zamora 2024 genera

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Ambulacraria

? Yanjiahella

Hemichordata

? Yanjiahella

Echinodermata

? Yanjiahella

Ctenoimbricata

Ctenocystoids†

Ctenocystis

Courtessolea

Cinctans†

Protocinctus

Solutes†

Coleicarpus

? Stylophorans†

Ceratocystis

Helicoplacoids†

Helicoplacus

Helicocystis

Eocrinoids†

Gogia

Edrioasteroids†

Camptostroma

Crinoids

Eleutherozoans
Echinozoans

Echinoidea

Holothuroidea

Asterozoa

Asteroidea

Ophiuroidea

Yanjiahella

[edit]

2019-02-15-a A stem group echinoderm from the basal Cambrian of China and the origins of Ambulacraria.pdf


Echinodermata

Yanjiahella

Ctenocystoids†

Cinctans†

Solutes†

Helicoplacus

Camptostroma

Eleutherozoans

Stromatocystites

Kinzercystis

Blastozoans†

Gogia

Akadocrinus

Pleurocystites

Ubaghsicystis

Macrocystella

Aethocrinus

  • Up to Helicoplacus is the stem group
  • Aethocrinus represents Crinozoans
  • Pelmatozoans was labled spanning Blastozoans and Crinozoans

Ridiculous chart

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Notable Linnaean taxonomies of phylum Echinodermata
Bather, 1900[69] Moore, 1966[70] Sprinkle, 1980[71]
  • Phylum Echinoderma [sic]
    • Grade Pelmatozoa
      •  
      •  
      • Class Cystidea†
      • Class Cystidea†
      • Class Cystidea†
      •  
      •  
      • Class Crinoidea
      • Class Cystidea†
      • Class Cystidea†
      • Class Cystidea†
      • Class Cystidea†
      •  
      • Class Blastoidea†
      •  
      •  
      •  
      •  
      • Class Edrioasteroidea†
      •  
      •  
      •  
      •  
    • Grade Eleutherozoa
      • Class Echinoidea
      • Class Holothuroidea
      •  
      • Class Stelleroidea
      • Class Stelleroidea†
  • Phylum Echinodermata
    •  
    • Subphylum Homalozoa†
      •  
      • Class Stylophora†
      • Class Homostelea†
      • Class Homoiostelea†
    • Subphylum Crinozoa
      • Class Paracrinoidea†
      • Class Crinoidea
      • Class Eocrinoidea†
      • Class Lepidocystoidea†
      • Class Cystoidea†
      • Class Cystoidea†
      • Class Parablastoidea†
      • Class Blastoidea†
      •  
      •  
      • Class Edrioblastoidea†
    • Subphylum Echinozoa
      • Class Edrioasteroidea†
      • Class Camptostromatoidea†
      • Class Helicoplacoidea†
      • Class Cyclocystoidea†
      • Class Ophiocistioidea†
      •  
      • Class Echinoidea
      • Class Holothuroidea
    • Subphylum Asterozoa
      • Class Stelleroidea
      • Class Stelleroidea†
  • Phylum Echinodermata
    •  
    • Subphylum Homalozoat
      • Class Ctenocystoidea†
      • Class Stylophora†
      • Class Homostelea†
      • Class Homoiostelea†
    • Subphylum Crinozoa
      • Class Paracrinoideat
      • Class Crinoidea
    • Subphylum Blastozoa†
      • Class Eocrinoidea†
      • Class Eocrinoidea†
      • Class Rhombifera†
      • Class Diploporita†
      • Class Parablastoideat
      • Class Blastoidea†
    • Subphylum Echinozoa
      • Class Edrioblastoidea†
      •  
      • Class Edrioasteroidea†
      • Class Edrioblasteroidea†
      • Class Helicoplacoideat
      • Class Cyclocystoidea†
      • Class Ophiocistioidea†
      •  
      • Class Echinoidea
      • Class Holothuroidea
    • Subphylum Asterozoa
      • Class Asteroidea
      • Class Ophiuroidea

Linnean Taxonomy Wide

[edit]
Bather, 1900 various authors, 1966-67 Ubaghs, 1978 Sprinkle, 1980
  • Phylum Echinoderma [sic]
    • Grade Pelmatozoa
      • Class Cystidea†
      • Class Blastoidea†
      • Class Crinoidea
      • Class Edrioasteroidea†
    • Grade Eleutherozoa
      • Class Holothuroidea
      • Class Stelleroidea
      • Class Echinoidea
  • Phylum Echinodermata
    • Subphylum Homalozoa†
      • Class Stylophora†
      • Class Homostelea†
      • Class Homoiostelea†
    • Subphylum Crinozoa
      • Class Lepidocystoidea†
      • Class Eocrinoidea†
      • Class Cystoidea†
      • Class Edrioblastoidea†
      • Class Parablastoidea†
      • Class Blastoidea†
      • Class Paracrinoidea†
      • Class Crinoidea
    • Subphylum Asterozoa
      • Class Stelleroidea
    • Subphylum Echinozoa
      • Class Helicoplacoidea†
      • Class Camptostromatoidea†
      • Class Edrioasteroidea†
      • Class Cyclocystoidea†
      • Class Ophiocystioidea†
      • Class Echinoidea
      • Class Holothuroidea
  • Phylum Echinodermata
    • Subphylum Homalozoa†
      • Class Ctenocystoidea†
      • Class Stylophora†
      • Class Homostelea†
      • Class Homoiostelea†
    • Subphylum Crinozoa
      • Class Eocrinoidea†
      • Class Rhombifera†
      • Class Diploporita†
      • Class Parablastoidea†
      • Class Blastoidea†
      • Class Paracrinoidea†
      • Class Crinoidea
    • Subphylum Asterozoa
      • Class Stelleroidea
    • Subphylum Echinozoa
      • Class Helicoplacoideat
      • Class Camptostromatoidea†
      • Class Edrioasteroidea†
      • Class Edrioblastoidea†
      • Class Cyclocystoidea†
      • Class Ophiocystioidea†
      • Class Echinoidea
      • Class Holothuroidea
  • Phylum Echinodermata
    • Subphylum Homalozoat
      • Class Ctenocystoidea†
      • Class Stylophora†
      • Class Homostelea†
      • Class Homoiostelea†
    • Subphylum Blastozoa†
      • Class Eocrinoidea†
      • Class Rhombifera†
      • Class Diploporita†
      • Class Parablastoideat
      • Class Blastoidea†
    • Subphylum Crinozoa
      • Class Paracrinoideat
      • Class Crinoidea
    • Subphylum Asterozoa
      • Class Asteroidea
      • Class Ophiuroidea
    • Subphylum Echinozoa
      • Class Helicoplacoideat
      • Class Edrioasteroidea†
      • Class Edrioblastoidea†
      • Class Cyclocystoidea†
      • Class Ophiocystioidea†
      • Class Echinoidea
      • Class Holothuroidea

Bather's Cystidea included taxa later assigned to Homalozoa, Eocrinoidea, and Cystoidea. The 1966-67 Cystoidea consisted of two orders, Rhombifera and Diplorita, that were promoted to classes in Sprinkle's phylogeny. Sprinkle split Blastozoa (taxa with brachioles) from Crinozoa (taxa with true arms), as well as demoting Lepidocystoidea and Camptostromatoidea, placing them under Eocrinoidea and Edrioasteroidea, respectively. MOAR NEEDED

Works cited

[edit]
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Potential specific citations

[edit]
  • Volume S phylogeny[72]
  • Volume S naming[73]
  • Tribrachidium heraldicum considered and dismisssed[74]
  • Arkarua adami rejected as true 5-fold symmetry is not seen in the Echinoderm fossil record until the emergence of Eleutherozoa in the Ordovician (or _possibly_ late Cambrian), and no other Echinoderm features are present[75]
  • Blastozoa to be paraphyletic with respect to Crinoidea[76]
  • Blastozoa (under the name Cystoidea) and Crinoidea to form a monophyletic slimmed-down Pelmatozoa[77]
  • Edrioasteroidea to be paraphyletic with respect to a clade consisting of Blastozoa and Crinoidea[48]
  • Rhombifera:Dichoporita:Glyptocystitoidea paraphyletic to Blastoidea, in polytomy with Rhombifera (Rhombiferdae), which is sister to Dichoporita:Hemicosmitoidea, which is sister to the rest of Glyptocystitoidea, with basal Glyptocystitida/Dichoporita:Sanducystis in polytome with it and several Eocrinoids[78]
  • Diploporita polyphyletic with Glyptosphaeritida paraphyletic w.r.t. Sphaeronitida, Rhombifera, Crinoidea, Paracrinoidea, Parablastoidea, and Diploporita:Asteroblastidae; Rhombifera paraphyletic to Coronoidea; Sphaeronitida monophyletic if includes former Glyptosphaeritidan Eucystis angelini; Holocystitidae monophyletic within Sphaeronitida[79]

Relationships

[edit]

Note that Smith (1984) uses "Cystoidea" in place of "Blastozoa", but makes the equivalence explicit.

Pelmatozoa Crinozoa Blastozoa Rhombifera Diploporita Coronoidea Lepidocystiodea Eocrinoidea Paracrinoidea Crinoidea Blastoidea Parablastoidea Edrioblastoidea Edrioasteroidea Stromatocystitoidea Camptostromatoidea Cyclocystoidea Ophiocistioidea Holothuroidea Echinoidea Echinozoa Ophiuroidea Stenuroidea Asteroidea Somasteroidea Asterozoa Eleutherozoa Helicocystiodea Helicoplacoidea Ctenoimbricatiodea Ctenocystoidea Cincta Soluta Stylophora Homalozoa
Pelmatozoa monophyletic: 1984[80] parent: 1984[80] parent: 1984[80] polytomy: 1984[80] polytomy: 1984[80]
Crinozoa
Blastozoa child: 1984[80]
Rhombifera sister: 2019[81]
Diploporita polyphyletic: 2019[79] paraphyletic: 2019[81]
Coronoidea sister: 2019[81]
Lepidocystoidea
Eocrinoidea
Paracrinoidea
Crinoidea child: 1984[80] excluded: 2019[81]
Blastoidea
Parablastoidea
Edrioblastoidea
Edrioasteroidea monophyletic: 1984[80] child: 1984[80] paraphyletic: 1984[80]
Stromatocystitoidea paraphyletic: 1984[80]
Camptostromatoidea polytomy: 1984[80] X polytomy: 1984[80]
Cyclocystoidea descendant: 1984[80] monophyletic: 1984[80]
Ophiocistioidea paraphyletic: 1984[80]
Holothuroidea stem: 1984[80]
Echinoidea
Echinozoa
Ophiuroidea
Stenuroidea
Asteroidea
Somasteroidea
Asterozoa
Eleutherozoa polytomy: 1984[80] scion: 1984[80] polytomy: 1984[80] monophyletic: 1984[80]
Helicocystoidea
Helicoplacoidea
Ctenoimbricatoidea
Ctenocystoidea
Cincta
Soluta
Stylophora
Homalozoa plesion: 1984[80]

References

[edit]
  1. ^ Edgecombe, Gregory D.; Giribet, Gonzalo; Dunn, Casey W.; Hejnol, Andreas; Kristensen, Reinhardt M.; Neves, Ricardo C.; Rouse, Greg W.; Worsaae, Katrine; Sørensen, Martin V. (June 2011). "Higher-level metazoan relationships: recent progress and remaining questions". Organisms, Diversity & Evolution. 11 (2): 151–172. Bibcode:2011ODivE..11..151E. doi:10.1007/s13127-011-0044-4. S2CID 32169826.
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