Orthonectida
Orthonectida (/ˌɔːrθəˈnɛktɪdə, -θoʊ-/[3]) is a small phylum of poorly known parasites of marine invertebrates[4] that are among the simplest of multi-cellular organisms. Members of this phylum are known as orthonectids.
| Orthonectida | |
|---|---|
| Two different female Orthonectids | |
| Scientific classification | |
| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Subkingdom: | Eumetazoa |
| Clade: | ParaHoxozoa |
| Clade: | Bilateria |
| Clade: | Nephrozoa |
| (unranked): | Protostomia |
| (unranked): | Spiralia |
| Clade: | Platytrochozoa |
| (unranked): | Mesozoa |
| Phylum: | Orthonectida Giard, 1877 [1][2] |
| Species | |
Biology edit
The adults, which are the sexual stage, are microscopic wormlike animals, consisting of a single layer of ciliated outer cells surrounding a mass of sex cells. They swim freely within the bodies of their hosts, which include flatworms, polychaete worms, bivalve molluscs, and echinoderms. Most are gonochoristic, with separate male and female individuals, but a few species are hermaphroditic.[5][6]
When they are ready to reproduce, adults leave the host, and sperm from the males penetrate the bodies of the females to achieve internal fertilisation. The resulting zygote develops into a ciliated larva that escapes from the mother to seek out new hosts. Once it finds a host, the larva loses its cilia and develops into a syncytial plasmodium larva. This, in turn, breaks up into numerous individual cells called agametes (ameiotic generative cells) which grow into the next generation of adults.[5][7]
Classification edit
The phylum consists of about 20 known species, of which Rhopalura ophiocomae is the best-known.[4] The phylum is not divided into classes or orders, and contains just two families.
Although originally described in 1877 as a class,[8] and later characterized as an order of the phylum Mesozoa, recent study has suggested that orthonectids are quite different from the rhombozoans, the other group in Mesozoa.[4] The genome of one orthonectid species, Intoshia linei, has been sequenced.[9] These animals are simplified spiralians. The genome data confirm earlier findings which allocated these organisms to Spiralia based on their morphology.[10]
Their position in the spiralian phylogenetic tree has yet to be determined. Some work appears to relate them to the Annelida[11][7] and, within the Annelida, finds them most closely allied to the Clitellata.[12] On the other hand, a 2022 study compensating for long-branch attraction has recovered the traditional grouping of Orthonectida with rhombozoans in a monophyletic Mesozoa placed close to Platyhelminthes or Gnathifera.[13] This supports a previous study which found orthonectids and rhombozoans to make a monophyletic taxon Mesozoa and form a clade with Rouphozoa (platyhelminths and gastrotrichs).[14]
Known species edit
Phylum Orthonectida
- Family Rhopaluridae Stunkard, 1937
- Ciliocincta
- Ciliocincta akkeshiensis Tajika, 1979 – Hokkaido, Japan; in flatworms (Turbellaria)
- Ciliocincta julini (Caullery and Mesnil, 1899) – E North Atlantic, in polychaetes
- Ciliocincta sabellariae Kozloff, 1965 – San Juan Islands, WA (USA); in polychaete (Neosabellaria cementarium)
- Intoshia
- Intoshia leptoplanae Giard, 1877 – E North Atlantic, in flatworms (Leptoplana)
- Intoshia linei Giard, 1877 – E North Atlantic, in nemertines (Lineus) = Rhopalura linei
- Intoshia major Shtein, 1953 – Arctic Ocean; in gastropods (Lepeta, Natica, Solariella) = Rhopalura major
- Intoshia metchnikovi (Caullery & Mesnil, 1899) – E North Atlantic, in polychaetes and nemertines
- Intoshia paraphanostomae (Westblad, 1942) – E North Atlantic, in flatworms (Acoela)
- Intoshia variabili (Alexandrov & Sljusarev, 1992) – Arctic Ocean, in flatworms (Macrorhynchus)
- Rhopalura
- Rhopalura elongata Shtein, 1953 – Arctic Ocean, in bivalves (Astarte)
- Rhopalura gigas (Giard, 1877)
- Rhopalura granosa Atkins, 1933 – E North Atlantic, in bivalves (Pododesmus)
- Rhopalura intoshi Metchnikoff – Mediterranean, in nemertines
- Rhopalura litoralis Shtein, 1953 – Arctic Ocean, in gastropods (Lepeta, Natica, Solariella)
- Rhopalura major Shtein, 1953
- Rhopalura murmanica Shtein, 1953 – Arctic Ocean, in gastropods (Rissoa, Columbella)
- Rhopalura ophiocomae Giard, 1877 – E North Atlantic, in ophiuroids (usually Amphipholis)
- Rhopalura pelseneeri Caullery & Mesnil, 1901 – E North Atlantic, polychaetes and nemertines
- Rhopalura philinae Lang, 1954 – E North Atlantic, in gastropods
- Rhopalura pterocirri de Saint-Joseph, 1896 – E North Atlantic, in polychaetes
- Rhopalura vermiculicola
- Stoecharthrum
- Stoecharthrum burresoni Kozloff, 1993
- Stoecharthrum fosterae Kozloff, 1993
- Stoecharthrum giardi Caullery & Mesnil, 1899 – E North Atlantic, in polychaetes
- Stoecharthrum monnati Kozloff, 1993 – E North Atlantic, in molluscs
- Ciliocincta
- Family Pelmatosphaeridae Stunkard, 1937
- Pelmatosphaera
- Pelmatosphaera polycirri Caullery and Mesnil, 1904 – E North Atlantic, in polychaetes and nemertines
- Pelmatosphaera
References edit
- ^ H. Furuya & J. van der Land (2010). "Orthonectida". World Register of Marine Species. Retrieved January 12, 2011.
- ^ "Orthonectida Giard, 1877". Integrated Taxonomic Information System. Retrieved January 12, 2011.
- ^ . Lexico UK English Dictionary. Oxford University Press. Archived from the original on 2020-03-22.
- ^ a b c Hanelt, B; Van Schyndel, D; Adema, C. M; Lewis, L. A; Loker, E. S (1996). "The phylogenetic position of Rhopalura ophiocomae (Orthonectida) based on 18S ribosomal DNA sequence analysis". Molecular Biology and Evolution. 13 (9): 1187–91. doi:10.1093/oxfordjournals.molbev.a025683. PMID 8896370.
- ^ a b Robert D. Barnes (1982). Invertebrate Zoology. Philadelphia, PA: Holt-Saunders International. pp. 247–248. ISBN 0-03-056747-5.
- ^ Sliusarev, G. S. (2003). "Orthonectida's life cycle". Parazitologiia. 37 (5): 418–427. PMID 14658313.
- ^ a b Zverkov, Oleg A.; Mikhailov, Kirill V.; Isaev, Sergey V.; Rusin, Leonid Y.; Popova, Olga V.; Logacheva, Maria D.; Penin, Alexey A.; Moroz, Leonid L.; Panchin, Yuri V.; Lyubetsky, Vassily A.; Aleoshin, Vladimir V. (24 May 2019). "Dicyemida and Orthonectida: Two Stories of Body Plan Simplification". Frontiers in Genetics. 10: 443. doi:10.3389/fgene.2019.00443. PMC 6543705. PMID 31178892.
- ^ Alfred Mathieu Giard (1877). "Sur les Orthonectida, classe nouvelle d'animaux parasites des Échinodermes et des Turbellariés" [On Orthonectida, a new class of parasitic animals of Echinoderms and Turbellarians]. Comptes Rendus (in French). 85 (18): 812–814.
- ^ Mikhailov, Kirill V; Slyusarev, Georgy S; Nikitin, Mikhail A; Logacheva, Maria D; Penin, Aleksey A; Aleoshin, Vladimir V; Panchin, Yuri V (2016). "The Genome of Intoshia linei Affirms Orthonectids as Highly Simplified Spiralians". Current Biology. 26 (13): 1768–74. doi:10.1016/j.cub.2016.05.007. PMID 27374341.
- ^ Sliusarev, G. S (2008). "Тип ортонектида (Orthonectida): строение, биология, положение в системе многоклеточных животных" [Phylum Orthonectida: Morphology, biology, and relationships to other multicellular animals]. Zhurnal Obshchei Biologii (in Russian). 69 (6): 403–27. PMID 19140332.
- ^ Bondarenko, N.; Bondarenko, A.; Starunov, V.; Slyusarev, G. (8 March 2019). "Comparative analysis of the mitochondrial genomes of Orthonectida: insights into the evolution of an invertebrate parasite species". Molecular Genetics and Genomics. 294 (3): 715–727. doi:10.1007/s00438-019-01543-1. PMID 30848356. S2CID 71716789.
- ^ Slyusarev, George S.; Starunov, Viktor V.; Bondarenko, Anton S.; Zorina, Natalia A.; Bondarenko, Natalya I. (April 2020). "Extreme Genome and Nervous System Streamlining in the Invertebrate Parasite Intoshia variabili". Current Biology. 30 (7): 1292–1298.e3. doi:10.1016/j.cub.2020.01.061. PMID 32084405.
- ^ Drábková, Marie; Kocot, Kevin M.; Halanych, Kenneth M.; Oakley, Todd H.; Moroz, Leonid L.; Cannon, Johanna T.; Kuris, Armand; Garcia-Vedrenne, Ana Elisa; Pankey, M. Sabrina; Ellis, Emily A.; Varney, Rebecca; Štefka, Jan; Zrzavý, Jan (6 July 2022). "Different phylogenomic methods support monophyly of enigmatic 'Mesozoa' (Dicyemida + Orthonectida, Lophotrochozoa)". Proceedings of the Royal Society B: Biological Sciences. 289 (1978): 20220683. doi:10.1098/rspb.2022.0683. PMC 9257288. PMID 35858055.
- ^ Lu, Tsai-Ming; Kanda, Miyuki; Satoh, Noriyuki; Furuya, Hidetaka (2017-05-29). "The phylogenetic position of dicyemid mesozoans offers insights into spiralian evolution". Zoological Letters. 3: 6. doi:10.1186/s40851-017-0068-5. ISSN 2056-306X. PMC 5447306. PMID 28560048.