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Rhizaria

The Rhizaria are an ill-defined but species-rich supergroup of mostly unicellular[1] eukaryotes.[2] Except for the Chlorarachniophytes and three species in the genus Paulinella in the phylum Cercozoa, they are all non-photosynthethic, but many foraminifera and radiolaria have a symbiotic relationship with unicellular algae.[3] A multicellular form, Guttulinopsis vulgaris, a cellular slime mold, has also been described.[4] This group was used by Cavalier-Smith in 2002, although the term "Rhizaria" had been long used for clades within the currently recognized taxon. Being described mainly from rDNA sequences, they vary considerably in form, having no clear morphological distinctive characters (synapomorphies), but for the most part they are amoeboids with filose, reticulose, or microtubule-supported pseudopods. In the absence of an apomorphy, the group is ill-defined, and its composition has been very fluid. Some Rhizaria possess mineral exoskeleton (thecae or loricas), which is in different clades within Rhizaria made out of opal (SiO2), celestite (SrSO4), or calcite (CaCO3). It can attain sizes of more than a centimeter with some species being able to form cylindrical colonies approximately 1 cm in diameter and greater than 1 m in length. They feed by capturing and engulfing prey with the extensions of their pseudopodia; forms that are symbiotic with unicellular algae contribute significantly to the total primary production of the ocean.[5]

Groups

The three main groups of Rhizaria are:[6]

A few other groups may be included in the Cercozoa, but some trees appear closer to the Foraminifera. These are the Phytomyxea and Ascetosporea, parasites of plants and animals, respectively, and the peculiar amoeba Gromia. The different groups of Rhizaria are considered close relatives based mainly on genetic similarities, and have been regarded as an extension of the Cercozoa. The name Rhizaria for the expanded group was introduced by Cavalier-Smith in 2002,[7] who also included the centrohelids and Apusozoa.

A noteworthy order that belongs to Ascetosporea is the Mikrocytida.[8] These are parasites of oysters. This includes the causative agent of Denman Island Disease, Mikrocytos mackini a small (2−3 μm diameter) amitochondriate protistan.[9]

Evolutionary relationships

Rhizaria are part of the SAR supergroup (Stramenopiles, Alveolates, Rhizaria), a grouping that had been presaged in 1993 through a study of mitochondrial morphologies.[10] SAR is currently placed in the Diaphoretickes along with Archaeplastida, Cryptista, Haptista, and several minor clades.

Historically, many rhizarians were considered animals because of their motility and heterotrophy. However, when a simple animal-plant dichotomy was superseded by a recognition of additional kingdoms, taxonomists generally placed amoebae in the kingdom Protista. When scientists began examining the evolutionary relationships among eukaryotes in the 1970's, it became clear that the kingdom Protista was paraphyletic. Rhizaria appear to share a common ancestor with Stramenopiles and Alveolates forming part of the SAR (Stramenopiles+Alveolates+Rhizaria) super assemblage.[11] Rhizaria has been supported by molecular phylogenetic studies as a monophyletic group.[12] Biosynthesis of 24-isopropyl cholestane precursors in various rhizaria[13] suggests a relevant ecological role already during the Ediacaran.

Phylogeny

Rhizaria is a monophyletic group composed of two sister phyla: Cercozoa and Retaria. Subsequently, Cercozoa and Retaria are also monophyletic.[14][15] The following cladogram depicts the evolutionary relationships between all rhizarian classes, and is made after the works of Cavalier-Smith et al. (2018),[16] Irwin et al. (2019)[17] and Sierra et al. (2022):[18]

Gallery

References

  1. ^ Christopher Taylor (2004). . Archived from the original on 2009-04-20.
  2. ^ Nikolaev SI, Berney C, Fahrni JF, et al. (May 2004). "The twilight of Heliozoa and rise of Rhizaria, an emerging supergroup of amoeboid eukaryotes". Proc. Natl. Acad. Sci. U.S.A. 101 (21): 8066–71. doi:10.1073/pnas.0308602101. PMC 419558. PMID 15148395.
  3. ^ Gast, Rebecca J.; Caron, David A. (2001-10-01). "Photosymbiotic associations in planktonic foraminifera and radiolaria". Hydrobiologia. 461 (1): 1–7. doi:10.1023/A:1012710909023. ISSN 1573-5117. S2CID 1387879.
  4. ^ Brown; et al. (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.
  5. ^ Caron, D. (2016). The rise of Rhizaria. Nature (London), 532(7600), 444–445. https://doi.org/10.1038/nature17892
  6. ^ Moreira D, von der Heyden S, Bass D, López-García P, Chao E, Cavalier-Smith T (July 2007). "Global eukaryote phylogeny: Combined small- and large-subunit ribosomal DNA trees support monophyly of Rhizaria, Retaria and Excavata". Mol. Phylogenet. Evol. 44 (1): 255–66. doi:10.1016/j.ympev.2006.11.001. PMID 17174576.
  7. ^ Cavalier-Smith, Thomas (2002). "The phagotrophic origin of eukaryotes and phylogenetic classification of Protozoa". International Journal of Systematic and Evolutionary Microbiology. 52 (2): 297–354. doi:10.1099/00207713-52-2-297. ISSN 1466-5026. PMID 11931142. Retrieved 2007-06-08.
  8. ^ Hartikainen, H; Stentiford, GD; Bateman, KS; Berney, C; Feist, SW; Longshaw, M; Okamura, B; Stone, D; Ward, G; Wood, C; Bass, D (2014). "Mikrocytids are a broadly distributed and divergent radiation of parasites in aquatic invertebrates" (PDF). Curr Biol. 24 (7): 807–12. doi:10.1016/j.cub.2014.02.033. PMID 24656829. S2CID 17180719.
  9. ^ Hine, Pm; Bower, Sm; Meyer, Gr; Cochennec-Laureau, N; Berthe, Fcj (2001). "Ultrastructure of Mikrocytos mackini, the cause of Denman Island disease in oysters Crassostrea spp. and Ostrea spp. in British Columbia, Canada". Diseases of Aquatic Organisms. 45 (3): 215–227. doi:10.3354/dao045215. ISSN 0177-5103. PMID 11558731.
  10. ^ Seravin LN. Osnovnye tipy i formy tonkogo stroeniia krist mitokhondriĭ: stepen' ikh évoliutsionnoĭ stabil'nosti (sposobnost' k morfologicheskim transformatsiiam) [The basic types and forms of the fine structure of mitochondrial cristae: the degree of their evolutionary stability (capacity for morphological transformations)]. Tsitologiia. 1993;35(4):3-34. Russian. PMID 8328023.
  11. ^ Burki, F; Shalchian-Tabrizi, K; Minge, M; Skjaeveland, A; Nikolaev, SI; Jakobsen, KS; Pawlowski, J (2007). Butler, Geraldine (ed.). "Phylogenomics Reshuffles the Eukaryotic Supergroups". PLoS ONE. 2 (8): e790–. Bibcode:2007PLoSO...2..790B. doi:10.1371/journal.pone.0000790. PMC 1949142. PMID 17726520.
  12. ^ Burki, Fabien; Shalchian-Tabrizi, Kamran; Pawlowski, Jan (August 23, 2008). "Phylogenomics reveals a new 'megagroup' including most photosynthetic eukaryotes". Biology Letters. 4 (4): 366–9. doi:10.1098/rsbl.2008.0224. PMC 2610160. PMID 18522922.
  13. ^ Hallmann, Christian; Stuhr, Marleen; Kucera, Michal; Zonneveld, Karin; Bobrovskiy, Ilya; Bowser, Samuel S.; Pawlowski, Jan; Deckker, Patrick De; Nowack, Eva C. M. (2019-03-04). "Putative sponge biomarkers in unicellular Rhizaria question an early rise of animals". Nature Ecology & Evolution. 3 (4): 577–581. doi:10.1038/s41559-019-0806-5. ISSN 2397-334X. PMID 30833757. S2CID 71148672.
  14. ^ Bass D, Chao EE, Nikolaev S, et al. (February 2009). "Phylogeny of Novel Naked Filose and Reticulose Cercozoa: Granofilosea cl. n. and Proteomyxidea Revised". Protist. 160 (1): 75–109. doi:10.1016/j.protis.2008.07.002. PMID 18952499.
  15. ^ Howe; et al. (2011), "Novel Cultured Protists Identify Deep-branching Environmental DNA Clades of Cercozoa: New Genera Tremula, Micrometopion, Minimassisteria, Nudifila, Peregrinia", Protist, 162 (2): 332–372, doi:10.1016/j.protis.2010.10.002, PMID 21295519
  16. ^ Cavalier-Smith, Thomas; Chao, Ema E.; Lewis, Rhodri (April 2018). "Multigene phylogeny and cell evolution of chromist infrakingdom Rhizaria: contrasting cell organisation of sister phyla Cercozoa and Retaria". Protoplasma. 255 (5): 1517–1574. doi:10.1007/s00709-018-1241-1. PMC 6133090. PMID 29666938.
  17. ^ Irwin, Nicholas A.T.; Tikhonenkov, Denis V.; Hehenberger, Elisabeth; Mylnikov, Alexander P.; Burki, Fabien; Keeling, Patrick J. (2019-01-01). "Phylogenomics supports the monophyly of the Cercozoa". Molecular Phylogenetics and Evolution. 130: 416–423. doi:10.1016/j.ympev.2018.09.004. ISSN 1055-7903. PMID 30318266. S2CID 52982396.
  18. ^ Sierra R, Mauffrey F, Cruz J, Holzmann M, Gooday AJ, Maurer-Alcalá X, Thakur R, Greco M, Weiner AKM, Katz LA, Pawlowski J (2022). "Taxon-rich transcriptomics supports higher-level phylogeny and major evolutionary trends in Foraminifera". Molecular Phylogenetics and Evolution. 174: 107546. doi:10.1016/j.ympev.2022.107546. ISSN 1055-7903. PMID 35690380.

External links

rhizaria, defined, species, rich, supergroup, mostly, unicellular, eukaryotes, except, chlorarachniophytes, three, species, genus, paulinella, phylum, cercozoa, they, photosynthethic, many, foraminifera, radiolaria, have, symbiotic, relationship, with, unicell. The Rhizaria are an ill defined but species rich supergroup of mostly unicellular 1 eukaryotes 2 Except for the Chlorarachniophytes and three species in the genus Paulinella in the phylum Cercozoa they are all non photosynthethic but many foraminifera and radiolaria have a symbiotic relationship with unicellular algae 3 A multicellular form Guttulinopsis vulgaris a cellular slime mold has also been described 4 This group was used by Cavalier Smith in 2002 although the term Rhizaria had been long used for clades within the currently recognized taxon Being described mainly from rDNA sequences they vary considerably in form having no clear morphological distinctive characters synapomorphies but for the most part they are amoeboids with filose reticulose or microtubule supported pseudopods In the absence of an apomorphy the group is ill defined and its composition has been very fluid Some Rhizaria possess mineral exoskeleton thecae or loricas which is in different clades within Rhizaria made out of opal SiO2 celestite SrSO4 or calcite CaCO3 It can attain sizes of more than a centimeter with some species being able to form cylindrical colonies approximately 1 cm in diameter and greater than 1 m in length They feed by capturing and engulfing prey with the extensions of their pseudopodia forms that are symbiotic with unicellular algae contribute significantly to the total primary production of the ocean 5 RhizariaTemporal range Neoproterozoic Recent Pha Proterozoic Archean Had nAmmonia tepida Foraminifera Scientific classificationDomain Eukaryota unranked DiaphoretickesKingdom ChromistaSubkingdom HarosaInfrakingdom RhizariaCavalier Smith 2002PhylaCercozoa Endomyxa Filosa Retaria Foraminifera Radiolaria Contents 1 Groups 2 Evolutionary relationships 2 1 Phylogeny 3 Gallery 4 References 5 External linksGroups EditFurther information wikispecies Rhizaria The three main groups of Rhizaria are 6 Cercozoa various amoebae and flagellates usually with filose pseudopods and common in soil Foraminifera amoeboids with reticulose pseudopods common as marine benthos Radiolaria amoeboids with axopods common as marine planktonA few other groups may be included in the Cercozoa but some trees appear closer to the Foraminifera These are the Phytomyxea and Ascetosporea parasites of plants and animals respectively and the peculiar amoeba Gromia The different groups of Rhizaria are considered close relatives based mainly on genetic similarities and have been regarded as an extension of the Cercozoa The name Rhizaria for the expanded group was introduced by Cavalier Smith in 2002 7 who also included the centrohelids and Apusozoa A noteworthy order that belongs to Ascetosporea is the Mikrocytida 8 These are parasites of oysters This includes the causative agent of Denman Island Disease Mikrocytos mackini a small 2 3 mm diameter amitochondriate protistan 9 Evolutionary relationships EditSee also Eukaryote Phylogeny Rhizaria are part of the SAR supergroup Stramenopiles Alveolates Rhizaria a grouping that had been presaged in 1993 through a study of mitochondrial morphologies 10 SAR is currently placed in the Diaphoretickes along with Archaeplastida Cryptista Haptista and several minor clades Historically many rhizarians were considered animals because of their motility and heterotrophy However when a simple animal plant dichotomy was superseded by a recognition of additional kingdoms taxonomists generally placed amoebae in the kingdom Protista When scientists began examining the evolutionary relationships among eukaryotes in the 1970 s it became clear that the kingdom Protista was paraphyletic Rhizaria appear to share a common ancestor with Stramenopiles and Alveolates forming part of the SAR Stramenopiles Alveolates Rhizaria super assemblage 11 Rhizaria has been supported by molecular phylogenetic studies as a monophyletic group 12 Biosynthesis of 24 isopropyl cholestane precursors in various rhizaria 13 suggests a relevant ecological role already during the Ediacaran Phylogeny Edit Rhizaria is a monophyletic group composed of two sister phyla Cercozoa and Retaria Subsequently Cercozoa and Retaria are also monophyletic 14 15 The following cladogram depicts the evolutionary relationships between all rhizarian classes and is made after the works of Cavalier Smith et al 2018 16 Irwin et al 2019 17 and Sierra et al 2022 18 Rhizaria Cercozoa Filosa Monadofilosa ImbricateaThecofiloseaSarcomonadeaHelkeseaMetromonadeaReticulofilosa GranofiloseaChlorarachneaEndomyxa PhytomyxeaGromiideaLapot guseviRetaria Foraminifera Globothalamea Monothalamea Tubothalamea Monothalamea PolycystineaAcanthareaSticholoncheaHalvaria HeterokontaAlveolataGallery Edit Cercomonas sp Cercozoa Cercomonadida Ebria sp Cercozoa Ebridea Rhipidodendron sp Cercozoa Spongomonadea Euglypha sp Cercozoa Euglyphida Phaeodarians Cercozoa Phaeodarea Clathrulina elegans Cercozoa Granofilosea Desmothoracida Chlorarachnion sp Cercozoa Chlorarachniophyta Vampyrella sp Cercozoa Vampyrellidae Gromia Cercozoa Gromiidea Powdery scab Cercozoa Plasmodiophorida Foraminiferans Retaria Foraminifera Polycystines Retaria Radiolaria Acantharians Retaria Radiolaria References Edit Christopher Taylor 2004 Rhizaria Archived from the original on 2009 04 20 Nikolaev SI Berney C Fahrni JF et al May 2004 The twilight of Heliozoa and rise of Rhizaria an emerging supergroup of amoeboid eukaryotes Proc Natl Acad Sci U S A 101 21 8066 71 doi 10 1073 pnas 0308602101 PMC 419558 PMID 15148395 Gast Rebecca J Caron David A 2001 10 01 Photosymbiotic associations in planktonic foraminifera and radiolaria Hydrobiologia 461 1 1 7 doi 10 1023 A 1012710909023 ISSN 1573 5117 S2CID 1387879 Brown et al 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 Caron D 2016 The rise of Rhizaria Nature London 532 7600 444 445 https doi org 10 1038 nature17892 Moreira D von der Heyden S Bass D Lopez Garcia P Chao E Cavalier Smith T July 2007 Global eukaryote phylogeny Combined small and large subunit ribosomal DNA trees support monophyly of Rhizaria Retaria and Excavata Mol Phylogenet Evol 44 1 255 66 doi 10 1016 j ympev 2006 11 001 PMID 17174576 Cavalier Smith Thomas 2002 The phagotrophic origin of eukaryotes and phylogenetic classification of Protozoa International Journal of Systematic and Evolutionary Microbiology 52 2 297 354 doi 10 1099 00207713 52 2 297 ISSN 1466 5026 PMID 11931142 Retrieved 2007 06 08 Hartikainen H Stentiford GD Bateman KS Berney C Feist SW Longshaw M Okamura B Stone D Ward G Wood C Bass D 2014 Mikrocytids are a broadly distributed and divergent radiation of parasites in aquatic invertebrates PDF Curr Biol 24 7 807 12 doi 10 1016 j cub 2014 02 033 PMID 24656829 S2CID 17180719 Hine Pm Bower Sm Meyer Gr Cochennec Laureau N Berthe Fcj 2001 Ultrastructure of Mikrocytos mackini the cause of Denman Island disease in oysters Crassostrea spp and Ostrea spp in British Columbia Canada Diseases of Aquatic Organisms 45 3 215 227 doi 10 3354 dao045215 ISSN 0177 5103 PMID 11558731 Seravin LN Osnovnye tipy i formy tonkogo stroeniia krist mitokhondriĭ stepen ikh evoliutsionnoĭ stabil nosti sposobnost k morfologicheskim transformatsiiam The basic types and forms of the fine structure of mitochondrial cristae the degree of their evolutionary stability capacity for morphological transformations Tsitologiia 1993 35 4 3 34 Russian PMID 8328023 Burki F Shalchian Tabrizi K Minge M Skjaeveland A Nikolaev SI Jakobsen KS Pawlowski J 2007 Butler Geraldine ed Phylogenomics Reshuffles the Eukaryotic Supergroups PLoS ONE 2 8 e790 Bibcode 2007PLoSO 2 790B doi 10 1371 journal pone 0000790 PMC 1949142 PMID 17726520 Burki Fabien Shalchian Tabrizi Kamran Pawlowski Jan August 23 2008 Phylogenomics reveals a new megagroup including most photosynthetic eukaryotes Biology Letters 4 4 366 9 doi 10 1098 rsbl 2008 0224 PMC 2610160 PMID 18522922 Hallmann Christian Stuhr Marleen Kucera Michal Zonneveld Karin Bobrovskiy Ilya Bowser Samuel S Pawlowski Jan Deckker Patrick De Nowack Eva C M 2019 03 04 Putative sponge biomarkers in unicellular Rhizaria question an early rise of animals Nature Ecology amp Evolution 3 4 577 581 doi 10 1038 s41559 019 0806 5 ISSN 2397 334X PMID 30833757 S2CID 71148672 Bass D Chao EE Nikolaev S et al February 2009 Phylogeny of Novel Naked Filose and Reticulose Cercozoa Granofilosea cl n and Proteomyxidea Revised Protist 160 1 75 109 doi 10 1016 j protis 2008 07 002 PMID 18952499 Howe et al 2011 Novel Cultured Protists Identify Deep branching Environmental DNA Clades of Cercozoa New Genera Tremula Micrometopion Minimassisteria Nudifila Peregrinia Protist 162 2 332 372 doi 10 1016 j protis 2010 10 002 PMID 21295519 Cavalier Smith Thomas Chao Ema E Lewis Rhodri April 2018 Multigene phylogeny and cell evolution of chromist infrakingdom Rhizaria contrasting cell organisation of sister phyla Cercozoa and Retaria Protoplasma 255 5 1517 1574 doi 10 1007 s00709 018 1241 1 PMC 6133090 PMID 29666938 Irwin Nicholas A T Tikhonenkov Denis V Hehenberger Elisabeth Mylnikov Alexander P Burki Fabien Keeling Patrick J 2019 01 01 Phylogenomics supports the monophyly of the Cercozoa Molecular Phylogenetics and Evolution 130 416 423 doi 10 1016 j ympev 2018 09 004 ISSN 1055 7903 PMID 30318266 S2CID 52982396 Sierra R Mauffrey F Cruz J Holzmann M Gooday AJ Maurer Alcala X Thakur R Greco M Weiner AKM Katz LA Pawlowski J 2022 Taxon rich transcriptomics supports higher level phylogeny and major evolutionary trends in Foraminifera Molecular Phylogenetics and Evolution 174 107546 doi 10 1016 j ympev 2022 107546 ISSN 1055 7903 PMID 35690380 External links EditMolecular Phylogeny of Amoeboid Protists Tree of Rhizaria Tree of Life Eukaryota https www nature com Retrieved from https en wikipedia org w index php title Rhizaria amp oldid 1134852943, wikipedia, wiki, book, books, library,

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