Orthonectida: Difference between revisions

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| image = Orthonetida_dict_flat_and_cylinder.png
| image = Orthonetida_dict_flat_and_cylinder.png
| image_caption = Two different female Orthonectids
| image_caption = Two different female Orthonectids
| display_parents = 8
| display_parents = 7
| taxon = Orthonectida  
| taxon = Orthonectida
| authority = [[Alfred Mathieu Giard|Giard]], 1877&nbsp;<ref>{{cite web |author1=H. Furuya  |author2=J. van der Land  |name-list-style=amp |year=2010 |title=Orthonectida |publisher=[[World Register of Marine Species]] |url=http://www.marinespecies.org/aphia.php?p=taxdetails&id=14220 |access-date=January 12, 2011}}</ref><ref>{{ITIS |id=57409 |taxon=Orthonectida Giard, 1877 |access-date=January 12, 2011}}</ref>
| authority = [[Alfred Mathieu Giard|Giard]], 1877&nbsp;<ref>{{cite web |author1=H. Furuya  |author2=J. van der Land  |name-list-style=amp |year=2010 |title=Orthonectida |publisher=[[World Register of Marine Species]] |url=http://www.marinespecies.org/aphia.php?p=taxdetails&id=14220 |access-date=January 12, 2011}}</ref><ref>{{ITIS |id=57409 |taxon=Orthonectida Giard, 1877 |access-date=January 12, 2011}}</ref>
| subdivision_ranks = Species
| subdivision_ranks = Species
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}}
}}


'''Orthonectida''' ({{IPAc-en|ˌ|ɔːr|θ|ə|ˈ|n|ɛ|k|t|ɪ|d|ə|,_|-|θ|oʊ|-}}{{refn|{{Cite dictionary |url=http://www.lexico.com/definition/Orthonectida |archive-url=https://web.archive.org/web/20200322182703/https://www.lexico.com/definition/orthonectida |url-status=dead |archive-date=2020-03-22 |title=Orthonectida |dictionary=[[Lexico]] UK English Dictionary |publisher=[[Oxford University Press]]}} }}) is a small [[phylum (biology)|phylum]] of poorly known [[parasite]]s of marine invertebrates<ref name="pmid8896370">{{cite journal |doi=10.1093/oxfordjournals.molbev.a025683 |pmid=8896370 |title=The phylogenetic position of Rhopalura ophiocomae (Orthonectida) based on 18S ribosomal DNA sequence analysis |journal=Molecular Biology and Evolution |volume=13 |issue=9 |pages=1187–91 |year=1996 |last1=Hanelt |first1=B |last2=Van Schyndel |first2=D |last3=Adema |first3=C. M |last4=Lewis |first4=L. A |last5=Loker |first5=E. S |doi-access=free }}</ref> that are among the simplest of multi-cellular organisms. Members of this phylum are known as '''orthonectids'''.
'''Orthonectida''' ({{IPAc-en|ˌ|ɔːr|θ|ə|ˈ|n|ɛ|k|t|ɪ|d|ə|,_|-|θ|oʊ|-}}{{refn|{{Cite dictionary |url=http://www.lexico.com/definition/Orthonectida |archive-url=https://web.archive.org/web/20200322182703/https://www.lexico.com/definition/orthonectida |archive-date=2020-03-22 |title=Orthonectida |dictionary=[[Lexico]] UK English Dictionary |publisher=[[Oxford University Press]]}} }}) is a small [[phylum (biology)|phylum]] of poorly known [[parasite]]s of marine invertebrates<ref name="pmid8896370">{{cite journal |doi=10.1093/oxfordjournals.molbev.a025683 |pmid=8896370 |title=The phylogenetic position of Rhopalura ophiocomae (Orthonectida) based on 18S ribosomal DNA sequence analysis |journal=Molecular Biology and Evolution |volume=13 |issue=9 |pages=1187–91 |year=1996 |last1=Hanelt |first1=B |last2=Van Schyndel |first2=D |last3=Adema |first3=C. M |last4=Lewis |first4=L. A |last5=Loker |first5=E. S |doi-access=free }}</ref> that are among the simplest of multi-cellular organisms. Members of this phylum are known as '''orthonectids'''.


==Biology==
==Biology==
The adults, which are the sexual stage, are microscopic wormlike animals, consisting of a single layer of [[cilia]]ted outer cells surrounding a mass of sex cells. They swim freely within the bodies of their hosts, which include [[flatworm]]s, [[polychaete]] worms, [[Bivalvia|bivalve]] molluscs, and [[echinoderm]]s. Most are [[gonochorism|gonochoristic]], with separate male and female individuals, but a few species are hermaphroditic.<ref name=IZ>{{cite book |author=Robert D. Barnes |year=1982 |title= Invertebrate Zoology |publisher= Holt-Saunders International |location= Philadelphia, PA|pages= 247–248|isbn= 0-03-056747-5}}</ref><ref>{{cite journal | url=https://pubmed.ncbi.nlm.nih.gov/14658313/ | pmid=14658313 | year=2003 | last1=Sliusarev | first1=G. S. | title=Orthonectida's life cycle | journal=Parazitologiia | volume=37 | issue=5 | pages=418–427 }}</ref>
The adults, which are the sexual stage, are microscopic wormlike animals, consisting of a single layer of [[cilia]]ted outer cells surrounding a mass of sex cells. They swim freely within the bodies of their hosts, which include [[flatworm]]s, [[polychaete]] worms, [[Bivalvia|bivalve]] molluscs, and [[echinoderm]]s. Most are [[gonochorism|gonochoristic]], with separate male and female individuals, but a few species are hermaphroditic.<ref name=IZ>{{cite book |author=Robert D. Barnes |year=1982 |title= Invertebrate Zoology |publisher= Holt-Saunders International |location= Philadelphia, PA|pages= 247–248|isbn= 0-03-056747-5}}</ref><ref>{{cite journal | pmid=14658313 | year=2003 | last1=Sliusarev | first1=G. S. | title=Orthonectida's life cycle | journal=Parazitologiia | volume=37 | issue=5 | pages=418–427 }}</ref>


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 [[syncytium|syncytial]] [[Plasmodium (life cycle)|plasmodium]] larva. This, in turn, breaks up into numerous individual cells called agametes (ameiotic generative cells) which grow into the next generation of adults.<ref name=IZ/><ref name=Zverkov2019>{{cite journal |last1=Zverkov |first1=Oleg A. |last2=Mikhailov |first2=Kirill V. |last3=Isaev |first3=Sergey V. |last4=Rusin |first4=Leonid Y. |last5=Popova |first5=Olga V. |last6=Logacheva |first6=Maria D. |last7=Penin |first7=Alexey A. |last8=Moroz |first8=Leonid L. |last9=Panchin |first9=Yuri V. |last10=Lyubetsky |first10=Vassily A. |last11=Aleoshin |first11=Vladimir V. |title=Dicyemida and Orthonectida: Two Stories of Body Plan Simplification |journal=Frontiers in Genetics |date=24 May 2019 |volume=10 |page=443 |doi=10.3389/fgene.2019.00443 |pmid=31178892 |pmc=6543705 |doi-access=free }}</ref>
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 [[syncytium|syncytial]] [[Plasmodium (life cycle)|plasmodium]] larva. This, in turn, breaks up into numerous individual cells called agametes (ameiotic generative cells) which grow into the next generation of adults.<ref name=IZ/><ref name=Zverkov2019>{{cite journal |last1=Zverkov |first1=Oleg A. |last2=Mikhailov |first2=Kirill V. |last3=Isaev |first3=Sergey V. |last4=Rusin |first4=Leonid Y. |last5=Popova |first5=Olga V. |last6=Logacheva |first6=Maria D. |last7=Penin |first7=Alexey A. |last8=Moroz |first8=Leonid L. |last9=Panchin |first9=Yuri V. |last10=Lyubetsky |first10=Vassily A. |last11=Aleoshin |first11=Vladimir V. |title=Dicyemida and Orthonectida: Two Stories of Body Plan Simplification |journal=Frontiers in Genetics |date=24 May 2019 |volume=10 |page=443 |doi=10.3389/fgene.2019.00443 |pmid=31178892 |pmc=6543705 |doi-access=free }}</ref>
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Although originally described in 1877 as a class,<ref>{{cite journal |author=Alfred Mathieu Giard |author-link=Alfred Mathieu Giard |year=1877 |title=Sur les Orthonectida, classe nouvelle d'animaux parasites des Échinodermes et des Turbellariés |trans-title=On Orthonectida, a new class of parasitic animals of Echinoderms and Turbellarians |journal=[[Comptes Rendus]] |volume=85 |issue=18 |pages=812–814 |language=fr |url=https://gallica.bnf.fr/ark:/12148/bpt6k30429/f808.item}}</ref> and later characterized as an order of the phylum [[Mesozoa]], a 1996 study has suggested that orthonectids are quite different from the [[rhombozoa]]ns, the other group in Mesozoa.<ref name="pmid8896370"/> The genome of one orthonectid species, ''[[Intoshia linei]]'', has been sequenced.<ref name=Mikhailov2016>{{cite journal |doi=10.1016/j.cub.2016.05.007 |pmid=27374341 |title=The Genome of Intoshia linei Affirms Orthonectids as Highly Simplified Spiralians |journal=Current Biology |volume=26 |issue=13 |pages=1768–74 |year=2016 |last1=Mikhailov |first1=Kirill V |last2=Slyusarev |first2=Georgy S |last3=Nikitin |first3=Mikhail A |last4=Logacheva |first4=Maria D |last5=Penin |first5=Aleksey A |last6=Aleoshin |first6=Vladimir V |last7=Panchin |first7=Yuri V |doi-access=free }}</ref> These animals are simplified [[spiralia]]ns. The genome data confirm earlier findings which allocated these organisms to Spiralia based on their morphology.<ref name=Sliusarev2008>{{cite journal |pmid=19140332 |year=2008 |last1=Sliusarev |first1=G. S |title=Тип ортонектида (Orthonectida): строение, биология, положение в системе многоклеточных животных |trans-title=Phylum Orthonectida: Morphology, biology, and relationships to other multicellular animals |language=ru |journal=Zhurnal Obshchei Biologii |volume=69 |issue=6 |pages=403–27 |url=https://elementy.ru/genbio/resume/193/Tip_ortonektida_Orthonectida_stroenie_biologiya_polozhenie_v_sisteme_mnogokletochnykh_zhivotnykh }}</ref>
Although originally described in 1877 as a class,<ref>{{cite journal |author=Alfred Mathieu Giard |author-link=Alfred Mathieu Giard |year=1877 |title=Sur les Orthonectida, classe nouvelle d'animaux parasites des Échinodermes et des Turbellariés |trans-title=On Orthonectida, a new class of parasitic animals of Echinoderms and Turbellarians |journal=[[Comptes Rendus]] |volume=85 |issue=18 |pages=812–814 |language=fr |url=https://gallica.bnf.fr/ark:/12148/bpt6k30429/f808.item}}</ref> and later characterized as an order of the phylum [[Mesozoa]], a 1996 study has suggested that orthonectids are quite different from the [[rhombozoa]]ns, the other group in Mesozoa.<ref name="pmid8896370"/> The genome of one orthonectid species, ''[[Intoshia linei]]'', has been sequenced.<ref name=Mikhailov2016>{{cite journal |doi=10.1016/j.cub.2016.05.007 |pmid=27374341 |title=The Genome of Intoshia linei Affirms Orthonectids as Highly Simplified Spiralians |journal=Current Biology |volume=26 |issue=13 |pages=1768–74 |year=2016 |last1=Mikhailov |first1=Kirill V |last2=Slyusarev |first2=Georgy S |last3=Nikitin |first3=Mikhail A |last4=Logacheva |first4=Maria D |last5=Penin |first5=Aleksey A |last6=Aleoshin |first6=Vladimir V |last7=Panchin |first7=Yuri V |doi-access=free }}</ref> These animals are simplified [[spiralia]]ns. The genome data confirm earlier findings which allocated these organisms to Spiralia based on their morphology.<ref name=Sliusarev2008>{{cite journal |pmid=19140332 |year=2008 |last1=Sliusarev |first1=G. S |title=Тип ортонектида (Orthonectida): строение, биология, положение в системе многоклеточных животных |trans-title=Phylum Orthonectida: Morphology, biology, and relationships to other multicellular animals |language=ru |journal=Zhurnal Obshchei Biologii |volume=69 |issue=6 |pages=403–27 |url=https://elementy.ru/genbio/resume/193/Tip_ortonektida_Orthonectida_stroenie_biologiya_polozhenie_v_sisteme_mnogokletochnykh_zhivotnykh }}</ref>


Their position in the spiralian phylogenetic tree has yet to be determined. Some work appears to relate them to the [[Annelida]]<ref name=Bondarenko2019>{{cite journal |last1=Bondarenko |first1=N. |last2=Bondarenko |first2=A. |last3=Starunov |first3=V. |last4=Slyusarev |first4=G. |title=Comparative analysis of the mitochondrial genomes of Orthonectida: insights into the evolution of an invertebrate parasite species |journal=Molecular Genetics and Genomics |date=8 March 2019 |volume=294 |issue=3 |pages=715–727 |doi=10.1007/s00438-019-01543-1 |pmid=30848356 |s2cid=71716789 }}</ref><ref name=Zverkov2019/> and, within the Annelida, finds them most closely allied to the [[Clitellata]].<ref name=Slyusarev2020>{{cite journal |last1=Slyusarev |first1=George S. |last2=Starunov |first2=Viktor V. |last3=Bondarenko |first3=Anton S. |last4=Zorina |first4=Natalia A. |last5=Bondarenko |first5=Natalya I. |title=Extreme Genome and Nervous System Streamlining in the Invertebrate Parasite Intoshia variabili |journal=Current Biology |date=April 2020 |volume=30 |issue=7 |pages=1292–1298.e3 |doi=10.1016/j.cub.2020.01.061 |pmid=32084405 |doi-access=free }}</ref> 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 (clade)|Gnathifera]].<ref>{{cite journal |last1=Drábková |first1=Marie |last2=Kocot |first2=Kevin M. |last3=Halanych |first3=Kenneth M. |last4=Oakley |first4=Todd H. |last5=Moroz |first5=Leonid L. |last6=Cannon |first6=Johanna T. |last7=Kuris |first7=Armand |last8=Garcia-Vedrenne |first8=Ana Elisa |last9=Pankey |first9=M. Sabrina |last10=Ellis |first10=Emily A. |last11=Varney |first11=Rebecca |last12=Štefka |first12=Jan |last13=Zrzavý |first13=Jan |title=Different phylogenomic methods support monophyly of enigmatic 'Mesozoa' (Dicyemida + Orthonectida, Lophotrochozoa) |journal=Proceedings of the Royal Society B: Biological Sciences |date=6 July 2022 |volume=289 |issue=1978 |pages=20220683 |doi=10.1098/rspb.2022.0683 |pmid=35858055 |pmc=9257288 }}</ref> This supports a previous study which found orthonectids and rhombozoans to make a monophyletic taxon Mesozoa and form a clade with [[Rouphozoa]] (platyhelminths and [[gastrotrich]]s).<ref>{{Cite journal|last1=Lu|first1=Tsai-Ming|last2=Kanda|first2=Miyuki|last3=Satoh|first3=Noriyuki|last4=Furuya|first4=Hidetaka|date=2017-05-29|title=The phylogenetic position of dicyemid mesozoans offers insights into spiralian evolution|journal=Zoological Letters|volume=3|pages=6|doi=10.1186/s40851-017-0068-5|issn=2056-306X|pmc=5447306|pmid=28560048 |doi-access=free }}</ref>
Their position in the spiralian phylogenetic tree has yet to be determined. Some work appears to relate them to the [[Annelida]]<ref name=Bondarenko2019>{{cite journal |last1=Bondarenko |first1=N. |last2=Bondarenko |first2=A. |last3=Starunov |first3=V. |last4=Slyusarev |first4=G. |title=Comparative analysis of the mitochondrial genomes of Orthonectida: insights into the evolution of an invertebrate parasite species |journal=Molecular Genetics and Genomics |date=8 March 2019 |volume=294 |issue=3 |pages=715–727 |doi=10.1007/s00438-019-01543-1 |pmid=30848356 |s2cid=71716789 }}</ref><ref name=Zverkov2019/> and, within the Annelida, finds them most closely allied to the [[Clitellata]].<ref name=Slyusarev2020>{{cite journal |last1=Slyusarev |first1=George S. |last2=Starunov |first2=Viktor V. |last3=Bondarenko |first3=Anton S. |last4=Zorina |first4=Natalia A. |last5=Bondarenko |first5=Natalya I. |title=Extreme Genome and Nervous System Streamlining in the Invertebrate Parasite Intoshia variabili |journal=Current Biology |date=April 2020 |volume=30 |issue=7 |pages=1292–1298.e3 |doi=10.1016/j.cub.2020.01.061 |pmid=32084405 |doi-access=free }}</ref> 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 (clade)|Gnathifera]].<ref>{{cite journal |last1=Drábková |first1=Marie |last2=Kocot |first2=Kevin M. |last3=Halanych |first3=Kenneth M. |last4=Oakley |first4=Todd H. |last5=Moroz |first5=Leonid L. |last6=Cannon |first6=Johanna T. |last7=Kuris |first7=Armand |last8=Garcia-Vedrenne |first8=Ana Elisa |last9=Pankey |first9=M. Sabrina |last10=Ellis |first10=Emily A. |last11=Varney |first11=Rebecca |last12=Štefka |first12=Jan |last13=Zrzavý |first13=Jan |title=Different phylogenomic methods support monophyly of enigmatic 'Mesozoa' (Dicyemida + Orthonectida, Lophotrochozoa) |journal=Proceedings of the Royal Society B: Biological Sciences |date=6 July 2022 |volume=289 |issue=1978 |article-number=20220683 |doi=10.1098/rspb.2022.0683 |pmid=35858055 |pmc=9257288 }}</ref> This supports a previous study which found orthonectids and rhombozoans to make a monophyletic taxon Mesozoa and form a clade with [[Rouphozoa]] (platyhelminths and [[gastrotrich]]s).<ref>{{Cite journal|last1=Lu|first1=Tsai-Ming|last2=Kanda|first2=Miyuki|last3=Satoh|first3=Noriyuki|last4=Furuya|first4=Hidetaka|date=2017-05-29|title=The phylogenetic position of dicyemid mesozoans offers insights into spiralian evolution|journal=Zoological Letters|volume=3|article-number=6|doi=10.1186/s40851-017-0068-5|issn=2056-306X|pmc=5447306|pmid=28560048 |doi-access=free }}</ref>


==Known species==
==Known species==
Line 40: Line 40:
*** ''[[Intoshia variabili]]'' <small>(Alexandrov & Sljusarev, 1992)</small> – Arctic Ocean, in flatworms (''[[Macrorhynchus]]'')
*** ''[[Intoshia variabili]]'' <small>(Alexandrov & Sljusarev, 1992)</small> – Arctic Ocean, in flatworms (''[[Macrorhynchus]]'')
** ''[[Rhopalura]]''  
** ''[[Rhopalura]]''  
*** ''[[Rhopalura elongata]]'' <small>Shtein, 1953</small> – Arctic Ocean, in bivalves (''[[Astarte]]'')
*** ''[[Rhopalura elongata]]'' <small>Shtein, 1953</small> – Arctic Ocean, in bivalves (''[[Astarte borealis|Astarte]]'')
*** ''[[Rhopalura gigas]]'' <small>(Giard, 1877)</small>
*** ''[[Rhopalura gigas]]'' <small>(Giard, 1877)</small>
*** ''[[Rhopalura granosa]]'' <small>Atkins, 1933</small> – E North Atlantic, in bivalves (''[[Pododesmus]]'')
*** ''[[Rhopalura granosa]]'' <small>Atkins, 1933</small> – E North Atlantic, in bivalves (''[[Pododesmus]]'')

Latest revision as of 23:35, 10 December 2025

Template:Short description Template:Automatic taxobox

Orthonectida (Template:IPAc-enTemplate:Refn) is a small phylum of poorly known parasites of marine invertebrates[1] that are among the simplest of multi-cellular organisms. Members of this phylum are known as orthonectids.

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.[2][3]

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.[2][4]

Classification

The phylum consists of about 20 known species, of which Rhopalura ophiocomae is the best-known.[1] The phylum is not divided into classes or orders, and contains just two families.

Although originally described in 1877 as a class,[5] 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.[1] The genome of one orthonectid species, Intoshia linei, has been sequenced.[6] These animals are simplified spiralians. The genome data confirm earlier findings which allocated these organisms to Spiralia based on their morphology.[7]

Their position in the spiralian phylogenetic tree has yet to be determined. Some work appears to relate them to the Annelida[8][4] and, within the Annelida, finds them most closely allied to the Clitellata.[9] 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.[10] 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).[11]

Known species

Phylum Orthonectida

References

Template:Sister project

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