Holothuroidea

 Specimen of Molpadia musculus, photographed by Carlo Fiori.

Belongs within: Echinozoa.
Contains: Elasipodida, Aspidochirotida, Dendrochirotacea, Apodida.

The Holothuroidea include the sea cucumbers, echinoderms in which the characteristic pentaradial symmetry of the phylum has been superimposed by a secondary bilaterality. Holothuroids have also reduced the skeletal plates to become much more flexible than other echinoderms. In members of the Apodacea, the tube feet have also been reduced or lost; mobility is by means of projecting anchor sclerites (Pawson 1966). The tentacles surrounding the mouth are modified in many holothuroid lineages; Apodacea retain simple digitate or pinnate tentacles but in Aspidochirotacea (Aspidochirotida and Elasipodida) they terminate in an approximately circular disc (Pawson 1966).

Characters (from Ubaghs 1978): Mostly free-living, usually with body elongated in oral-aboral axis and secondary bilateral symmerry; mouth encircled by tentacles; pharynx surrounded· by calcareous ring; ambulacral grooves lacking; skeleton ordinarily consisting of microscopic sclerites embedded in body wall; gonad single.

<==Holothuroidea (see below for synonymy)
    |--AspidochirotaceaU78
    |    |--ElasipodidaS97
    |    `--+--AspidochirotidaS97
    |       `--DendrochirotaceaS97
    `--ApodaceaU78
         |--ApodidaS97
         `--MolpadiidaU78
              |  i. s.: Cherbonniera utriculus Sibuet 1974B01
              |         Calcligula Frizzell & Exline 1956P66, FE66 [incl. Molpadioites Deflandre-Rigaud 1961FE66]
              |           |--*C. perforata Frizzell & Exline 1956FE66
              |           |--C. elgeriFE66
              |           |--C. fictaFE66
              |           `--C. jurassicaFE66
              |--GephyrothuriidaeP66
              |--CaudinidaeMG-H11
              |    |--Acaudina leucoprocta (Clark 1938)MG-H11
              |    |--Caudina coriaceaH04
              |    |--Pedatopriscus Deflandre-Rigaud 1961SG93, FE66
              |    |    `--*P. pinguis (Deflandre-Rigaud 1946) [=Priscopedatus pinguis]FE66
              |    |--AphelodactylaMG-H11
              |    |    |--A. bacilla (Cherbonnier & Féral 1981)MG-H11
              |    |    |--A. molpadioides (Semper 1868)MG-H11
              |    |    `--A. suspecta (Cherbonnier & Féral 1981)MG-H11
              |    `--Paracaudina chilensis (Muller 1850)SM15
              `--MolpadiidaeMG-H11
                   |--Heteromolpadia tridens (Sluiter 1901)MG-H11
                   |--Trochostoma Danielsen & Koren 1878MG-H11, B01
                   |    `--T. parvulum Cherbonnier & Féral 1981MG-H11
                   |--Staurocumites Deflandre-Rigaud 1952SG93, FE66
                   |    `--*S. bartensteini Deflandre-Rigaud 1952FE66
                   |--Calcancorella Deflandre-Rigaud 1961SG93, FE66 [Calcancorellidae]
                   |    `--*C. spectabilis (Deflandre-Rigaud 1959) [=Synaptites (Calcancora) spectabilis]FE66
                   `--MolpadiaMG-H11
                        |--M. andamanensis (Walsh 1981)MG-H11
                        |--M. musculus Risso 1826MG-H11
                        `--M. ooliticaBK77
Holothuroidea incertae sedis:
  Stereoderma unisemitaK-M02
  Pseudocucumis mixtaW79
  Protancyra challengeriJ18
  Patinapta ooplax (Marenzeller 1881)S-PR86
  Ocellothuria Boczarowski 2001B01
    `--*O. binoculata Boczarowski 2001B01
  Canisia Mostler 1972B01
  Triradites Mostler 1969B01
  Pseudothyone sculponea Cherbonnier 1958B01
  Andenothyone gondwanensis Haude 1995B01
  Nudicorona seilacheri Haude 1995B01
  Bathyplotes natansBK77
  Palaeocucumariidae [Arthrochirotida, Etheridgellidae]B01
    |--Frizzellus Hampton 1958FE66
    |    `--*F. irregularis Hampton 1958FE66
    |--Etheridgella Croneis 1932B01, FE66
    |    |--*E. porosa Croneis 1932FE66
    |    `--E. biconvexaFE66
    `--Palaeocucumaria Lehmann 1958B01
         |--*P. hunsrueckiana Lehmann 1958B01
         |--P. ancile Boczarowski 2001B01
         `--P. delicata Boczarowski 2001B01
  CucumeriaMW07
    |--C. salvaMW07
    `--C. sykionMW07
  Kaliobullites Kristan-Tollmann 1963 [Kaliobullitidae]FE66
    `--*K. umbo Kristan-Tollmann 1963FE66
  Alexandrites Kristan-Tollmann 1964 [Alexandritidae]FE66
    `--*A. alexandri Kristan-Tollmann 1964FE66
  Oerstergrenia digitataPP64
  TrachythyonePP64
    |--T. elongataPP64
    `--T. tergestinaPP64
  Punctatites [Punctatitidae]SG93
    |--P. aequiperforatus Mostler & Rahimi-Yazd 1976SG93
    `--P. triplex Mostler 1972SG93
  Kozurella [Kozurellidae]SG93
    `--K. formosa Mostler 1972SG93
  Pentacta doliolumG20
  Pentadactyla longidentisPG98

Holothuroidea [Ascidiastella, Cylindroides, Fistulidae, Fistulides, Holothuriacea, Holothuriadae, Holothuriae, Holothurida, Holothuridae, Holothuridea, Holothurina, Holothurioidea, Holothuromorpha, Scytactinata, Scytodermata]

*Type species of generic name indicated

References

[BK77] Barel, C. D. N., & P. G. N. Kramers. 1977. A survey of the echinoderm associates of the north-east Atlantic area. Zoologische Verhandelingen 156: 1–159.

[B01] Boczarowski, A. 2001. Isolated sclerites of Devonian non-pelmatozoan echinoderms. Palaeontologia Polonica 59: 1–219.

[FE66] Frizzell, D. L., & H. Exline. 1966. Holothuroidea—fossil record. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt U. Echinodermata 3 vol. 2 pp. U646–U672. The Geological Society of America, Inc., and The University of Kansas Press.

[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.

[H04] Haeckel, E. 1899–1904. Kunstformen der Natur. Bibliographisches Institut: Leipzig und Wien.

[J18] Jaekel, O. 1918. Phylogenie und System der Pelmatozoen. Paläontologische Zeitschrift 3: 1–128.

[K-M02] Klein-MacPhee, G. 2002. Righteye flounders. Family Pleuronectidae. In: Collette, B. B., & G. Klein-MacPhee (eds) Bigelow and Schroeder’s Fishes of the Gulf of Maine 3rd ed. pp. 560–587. Smithsonian Institute Press: Washington.

[MW07] Mallatt, J., & C. J. Winchell. 2007. Ribosomal RNA genes and deuterostome phylogeny revisited: more cyclostomes, elasmobranchs, reptiles, and a brittle star. Molecular Phylogenetics and Evolution 43: 1005–1022.

[MG-H11] McEnnulty, F. R., K. L. Gowlett-Holmes, A. Williams, F. Althaus, J. Fromont, G. C. B. Poore, T. D. O’Hara, L. Marsh, P. Kott, S. Slack-Smith, P. Alderslade & M. V. Kitahara. 2011. The deepwater megabenthic invertebrates on the western continental margin of Australia (100–1100 m depths): composition, distribution and novelty. Records of the Western Australian Museum Supplement 80: 1–191.

[P66] Pawson, D. L. 1966. Phylogeny and evolution of holothuroids. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt U. Echinodermata 3 vol. 2 pp. U641–U646. The Geological Society of America, Inc., and The University of Kansas Press.

[PP64] Peres, J. M., & J. Picard. 1964. Nouveau manuel de bionomie benthique de la mer Mediterranee. Recueil des Travaux de la Station Marine d’Endoume, Bulletin 31 (27): 5–137.

[PG98] Probert, P. K., & S. L. Grove. 1998. Macrobenthic assemblages of the continental shelf and upper slope off the west coast of South Island, New Zealand. Journal of the Royal Society of New Zealand 28: 259–280.

[S-PR86] Salvini-Plawen, L. v., & G. C. Rao. 1986. Echinoderma. In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 701–704. E. J. Brill/Dr W. Backhuys: Leiden.

[SM15] Sampey, A., & L. M. Marsh. 2015. Kimberley marine biota. Historical data: echinoderms. Records of the Western Australian Museum Supplement 84: 207–246.

[SG93] Simms, M. J., A. S. Gale, P. Gilliland, E. P. F. Rose & G. D. Sevastopulo. 1993. Echinodermata. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 491–528. Chapman & Hall: London.

[S97] Smith, A. B. 1997. Echinoderm larvae and phylogeny. Annual Review of Ecology and Systematics 28: 219–241.

[U78] Ubaghs, G. 1978. Classification of the echinoderms. In: Moore, R. C., & C. Teichert (eds) Treatise on Invertebrate Paleontology pt T. Echinodermata 2. Crinoidea vol. 1 pp. T359–T367. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas: Lawrence (Kansas).

[W79] Warner, G. F. 1979. Aggregation in echinoderms. In: Larwood, G., & B. R. Rosen (eds) Biology and Systematics of Colonial Organisms pp. 375–396. Academic Press: London.

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