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Helkesida

From Wikipedia, the free encyclopedia
Group of eukaryotic microorganisms

Helkesida
Scientific classificationEdit this classification
Domain:Eukaryota
Clade:Diaphoretickes
Clade:SAR
Clade:Rhizaria
Phylum:Cercozoa
Superclass:Eoglissa
Class:Helkesea
Cavalier-Smith 2018
Order:Helkesida
Cavalier-Smith 2018[1]
Families[1][2]
Diversity
24 species

Helkesida (formerly known asSainouroidea)[1] is a group ofmicroscopicprotists belonging to thesupergroupRhizaria, both discovered throughmolecular phylogenetic analyses. It containsamoeboidflagellates with twoflagella. They are either free-living, mostly onfecal matter, or live inside thegut of animals. Among these amoebae, one lineage has independently evolved aggregativemulticellularity similarly toslime moulds.[3]

Biology

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The organisms classified as Helkesida commonly have agliding motility in which thecells glide on their posteriorflagellum. They are ancestrallyamoeboid bi-flagellates without scales ortheca.[2] Unlike mostCercozoa which have tubularmitochondrial cristae, they can also present flat cristae or discoid cristae. They are the only group withinRhizaria that present discoid mitochondrial cristae.[4]

These organisms have an amorphous apicalcentrosome attached to the nucleus by arhizoplast. Thekinetid arises from 2–4 very shortcentrioles with dense fibrous roots that attach them to each other and to the nucleus. Their anterior flagellum is reduced to a stub without its 9+2axoneme. The centrosome also generates numerousmicrotubules in larger cells. TheGolgi apparatus is seen attached to thenuclear envelope and the anterior rhizoplast. They have a microbody attached to the posterior end of thenucleus.[5]

One helkesid genus,Guttulinopsis, represents an independent lineage in which aggregativemulticellularity hasevolved to generate "fungi-like" fruiting bodies calledsorocarps, similarly toslime moulds such asDictyostelium.[4]

Ecology

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The helkesidamoebae arebacterivores that can be free-living, mostly associated tofecal environments, orendozoic, associated toanimals.[6] They thrive inaerobic conditions and themicroaerophilicgut environment of animals.Rosculus can thrive inanaerobic culture. It is unknown if their preferred habitat is free-living or endozoic.[3]

Some host species can harbor different helkesid genera and species. One animal can be infected by multiple species simultaneously, and one species can also infect different animal hosts. More sampling of hosts, amoebae and molecular data is needed to better understand the life history and ecology of these protists.[3]

Evolution and systematics

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History

[edit]

Helkesida is a group initially named Sainouroidea. It was discovered in 2009 as a highly divergent clade withinCercozoa throughphylogenetic analyses that used thesequencing of18S ribosomal RNA fromCholamonas cytrodiopsidis,Sainouron acronematica andHelkesimastix marina. It is a molecularly diverse clade that branches within a group of ancestrallyamoeboid bi-flagellates that usually lack an outer cell coat, known asMonadofilosa.[5] A 2016 study revealed a previously unknown wide diversity of Sainouroidea in fecal environments. Previousenvironmental samplings excluded sequences from Sainouroidea due to their highly divergent 18S rDNA sequences.[6] A 2018 study described several new genera and species.[3]

The initial name for this group, Sainouroidea, had the-oidea suffix for superfamilies, but it was not assigned to any existing classes or orders due to the uncertainty of its phylogenetic position.[5] In a 2018 revision, the classHelkesea and orderHelkesida were created as a substitute for this name. Sainouroidea was then modified to only include one of the three helkesid families,Sainouridae. A second superfamily,Helkesimastigoidea, was created to host the remaining two families,Helkesimastigidae andGuttulinopsidae.[1]

Classification

[edit]
Phylogeny of Helkesida
Cladogram of Helkesida, based on a18S rDNA phylogenetic analysis within a 2018 study.[3] The study names this group "Sainouroidea" due to being published prior to the taxonomic change to Helkesida.[1]

Currently, Helkesida contains 24species distributed in 9genera, 3families[2] and 2superfamilies.[1] Additionally, manyOTUs found throughenvironmental sequencing may represent undescribed clades.[4][6]

References

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  1. ^abcdefThomas Cavalier-Smith; Ema E-Yung Chao; Rhodri Lewis (17 April 2018)."Multigene phylogeny and cell evolution of chromist infrakingdom Rhizaria: contrasting cel organisation of sister phyla Cercozoa and Retaria".Protoplasma.doi:10.1007/S00709-018-1241-1.ISSN 0033-183X.PMC 6133090.PMID 29666938.Wikidata Q53073157.
  2. ^abcAdl SM, Bass D, Lane CE, Lukeš J, Schoch CL, Smirnov A, Agatha S, Berney C, Brown MW, Burki F, Cárdenas P, Čepička I, Chistyakova L, del Campo J, Dunthorn M, Edvardsen B, Eglit Y, Guillou L, Hampl V, Heiss AA, Hoppenrath M, James TY, Karnkowska A, Karpov S, Kim E, Kolisko M, Kudryavtsev A, Lahr DJG, Lara E, Le Gall L, Lynn DH, Mann DG, Massana R, Mitchell EAD, Morrow C, Park JS, Pawlowski JW, Powell MJ, Richter DJ, Rueckert S, Shadwick L, Shimano S, Spiegel FW, Torruella G, Youssef N, Zlatogursky V, Zhang Q (2019)."Revisions to the Classification, Nomenclature, and Diversity of Eukaryotes".Journal of Eukaryotic Microbiology.66 (1):4–119.doi:10.1111/jeu.12691.PMC 6492006.PMID 30257078.
  3. ^abcdefghiSchuler GA, Tice AK, Pearce RA, Foreman E, Stone J, Gammill S, Willson JD, Reading C, Silberman JD, Brown MW (2018)."Phylogeny and Classification of Novel Diversity in Sainouroidea (Cercozoa, Rhizaria) Sheds Light on a Highly Diverse and Divergent Clade"(PDF).Protist.169 (6):853–874.doi:10.1016/j.protis.2018.08.002.PMID 30415103.S2CID 53289638.
  4. ^abcBrown MW, Kolisko M, Silberman JD, Roger AJ (2012)."Aggregative Multicellularity Evolved Independently in the Eukaryotic Supergroup Rhizaria".Current Biology.22 (12):1123–1127.doi:10.1016/j.cub.2012.04.021.PMID 22608512.S2CID 17510471.
  5. ^abcdCavalier-Smith T, Lewis R, Chao EE, Oates B, Bass D (2009). "Helkesimastix marina n. sp. (Cercozoa: Sainouroidea superfam. n.) a Gliding Zooflagellate of Novel Ultrastructure and Unusual Ciliary Behaviour".Protist.160 (3):452–479.doi:10.1016/j.protis.2009.03.003.PMID 19523874.
  6. ^abcBass D, Silberman JD, Brown MW, Pearce RA, Tice AK, Jousset A, Geisen S, Hartikainen H (2016). "Coprophilic amoebae and flagellates, including Guttulinopsis, Rosculus and Helkesimastix, characterise a divergent and diverse rhizarian radiation and contribute to a large diversity of faecal-associated protists".Environ Microbiol.18 (5):1604–1619.doi:10.1111/1462-2920.13235.PMID 26914587.
  7. ^Flavin M, O'Kelly CJ, Nerad TA, Wilkinson G (2000). "Cholamonas cyrtodiopsidis gen. n., sp. n.(Cercomonadida), an endocommensal, mycophagous heterotrophic flagellate with doubled kinetid".Acta Protozoologica.39 (1):51–60.
  8. ^Sandon H (1924). "Some Protozoa from the Soils and Mosses of Spitsbergen. Results of the Oxford University Expedition to Spitsbergen, No. 27".Zoological Journal of the Linnean Society.35 (237):449–475.doi:10.1111/j.1096-3642.1924.tb00051.x.
  9. ^Cavalier-Smith T, Lewis R, Chao EE, Oates B, Bass D (2008). "Morphology and Phylogeny of Sainouron acronematica sp. n. and the Ultrastructural Unity of Cercozoa".Protist.159 (4):591–620.doi:10.1016/j.protis.2008.04.002.PMID 18583188.
  10. ^Woodcock HM, Lapage G (1915). "Observations on the life-cycle of a new flagellate, Helkesimastix fæcicola, n. g., n. sp.: Together remarks on the question of syngamy in the trypanosomes".Proc. R. Soc. Lond. B.88 (604):353–370.doi:10.1098/rspb.1915.0001.
  11. ^Olive EW (1901)."A Preliminary Enumeration of the Sorophoreæ".Proceedings of the American Academy of Arts and Sciences.37 (12):333–344.doi:10.2307/20021671.JSTOR 20021671.
  12. ^Hawes RS (1963). "On Rosculus ithacus gen. n., sp. n. (Protozoa, Amoebina), with special reference to its mitosis and phylogenetic relations".Journal of Morphology.113 (2):139–149.doi:10.1002/jmor.1051130202.PMID 14061990.S2CID 40212732.
Rhizaria classification
 Cercozoa 
Reticulofilosa*
Skiomonadea
Granofilosea
Chlorarachneaa
Monadofilosa
Eoglissa
Metromonadea
Helkeseaaf
Ventrifilosa
Sarcomonadea*af
Imbricatea
Silicofilosea
Thecofilosea
Endomyxa
Marimyxia
Gromiidea
Ascetosporeas
Proteomyxia
   Retaria   
Radiolariah
Sticholonchea
Acantharea
Polycystinea
Foraminiferaa
Monothalamea*
Tubothalamea
Globothalamea
Incertae sedis
Incertae sedis
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