Orthonectida

Biology

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.

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.

Classification

The phylum consists of about 20 known species, of which Rhopalura ophiocomae is the best-known. The phylum is not divided into classes or orders, and contains just two families.

Although originally described in 1877 as a class, and later characterized as an order of the phylum Mesozoa, a 1996 study has suggested that orthonectids are quite different from the rhombozoans, the other group in Mesozoa. The genome of one orthonectid species, Intoshia linei, has been sequenced. These animals are simplified spiralians. The genome data confirm earlier findings which allocated these organisms to Spiralia based on their morphology.

Their position in the spiralian phylogenetic tree has yet to be determined. Some work appears to relate them to the Annelida 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. 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).

Known species

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
• Family Pelmatosphaeridae Stunkard, 1937
  • Pelmatosphaera
    • Pelmatosphaera polycirri Caullery and Mesnil, 1904 – E North Atlantic, in polychaetes and nemertines

References

References

1. (2010). "Orthonectida". [[World Register of Marine Species]].
2. {{ITIS
3. (1996). "The phylogenetic position of Rhopalura ophiocomae (Orthonectida) based on 18S ribosomal DNA sequence analysis". Molecular Biology and Evolution.
4. Robert D. Barnes. (1982). "Invertebrate Zoology". Holt-Saunders International.
5. (2003). "Orthonectida's life cycle". Parazitologiia.
6. (24 May 2019). "Dicyemida and Orthonectida: Two Stories of Body Plan Simplification". Frontiers in Genetics.
7. Alfred Mathieu Giard. (1877). "Sur les Orthonectida, classe nouvelle d'animaux parasites des Échinodermes et des Turbellariés". [[Comptes Rendus]].
8. (2016). "The Genome of Intoshia linei Affirms Orthonectids as Highly Simplified Spiralians". Current Biology.
9. (2008). "Тип ортонектида (Orthonectida): строение, биология, положение в системе многоклеточных животных". Zhurnal Obshchei Biologii.
10. (8 March 2019). "Comparative analysis of the mitochondrial genomes of Orthonectida: insights into the evolution of an invertebrate parasite species". Molecular Genetics and Genomics.
11. (April 2020). "Extreme Genome and Nervous System Streamlining in the Invertebrate Parasite Intoshia variabili". Current Biology.
12. (6 July 2022). "Different phylogenomic methods support monophyly of enigmatic 'Mesozoa' (Dicyemida + Orthonectida, Lophotrochozoa)". Proceedings of the Royal Society B: Biological Sciences.
13. (2017-05-29). "The phylogenetic position of dicyemid mesozoans offers insights into spiralian evolution". Zoological Letters.