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The '''Rhizaria''' are a diverse and species-rich supergroup of mostly ]<ref>{{cite web |url=http://www.palaeos.com/Eukarya/Units/Rhizaria/Rhizaria.html |year=2004 |last=Taylor |first=Christopher |title=Rhizaria |url-status=dead |archive-url=https://web.archive.org/web/20090420034036/http://www.palaeos.com/Eukarya/Units/Rhizaria/Rhizaria.html |archive-date=2009-04-20 }}</ref> ]s.<ref name="pmid15148395">{{cite journal |last=Nikolaev |first=Sergey I. |last2=Berney |first2=Cédric |last3=Fahrni |first3=José F. |last4=Bolivar |first4=Ignacio |last5=Polet |first5=Stephane |last6=Mylnikov |first6=Alexander P. |last7=Aleshin |first7=Vladimir V. |last8=Petrov |first8=Nikolai B. |last9=Pawlowski |first9=Jan |display-authors=3 |title=The twilight of Heliozoa and rise of Rhizaria, an emerging supergroup of amoeboid eukaryotes |journal=PNAS |volume=101 |issue=21 |pages=8066–71 |date=May 2004 |pmid=15148395 |pmc=419558 |doi=10.1073/pnas.0308602101 |doi-access=free }}</ref> Except for the ] and three species in the genus ] in the phylum ], they are all non-photosynthethic, but many foraminifera and radiolaria have a symbiotic relationship with unicellular algae.<ref>{{Cite journal |last1=Gast |first1=Rebecca J.|last2=Caron |first2=David A. |date=2001-10-01 |title=Photosymbiotic associations in planktonic foraminifera and radiolaria |journal=Hydrobiologia |volume=461 |issue=1 |pages=1–7 |doi=10.1023/A:1012710909023 |s2cid=1387879}}</ref> A multicellular form, ''Guttulinopsis vulgaris'', a cellular ], has been described.<ref>{{cite journal |last=Brown |first=Matthew W. |last2=Kolisko |first2=Martin |last3=Silberman |first3=Jeffrey D. |last4=Roger |first4=Andrew J. |title=Aggregative Multicellularity Evolved Independently in the Eukaryotic Supergroup Rhizaria |journal=] |volume=22 |issue=12 |year=2012 |doi=10.1016/j.cub.2012.04.021 |pages=1123–1127|doi-access=free }}</ref> |
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The '''Rhizaria''' are a diverse and species-rich supergroup of mostly ]<ref>{{cite web |url=http://www.palaeos.com/Eukarya/Units/Rhizaria/Rhizaria.html |year=2004 |last=Taylor |first=Christopher |title=Rhizaria |url-status=dead |archive-url=https://web.archive.org/web/20090420034036/http://www.palaeos.com/Eukarya/Units/Rhizaria/Rhizaria.html |archive-date=2009-04-20 }}</ref> ]s.<ref name="pmid15148395">{{cite journal |last=Nikolaev |first=Sergey I. |last2=Berney |first2=Cédric |last3=Fahrni |first3=José F. |last4=Bolivar |first4=Ignacio |last5=Polet |first5=Stephane |last6=Mylnikov |first6=Alexander P. |last7=Aleshin |first7=Vladimir V. |last8=Petrov |first8=Nikolai B. |last9=Pawlowski |first9=Jan |display-authors=3 |title=The twilight of Heliozoa and rise of Rhizaria, an emerging supergroup of amoeboid eukaryotes |journal=PNAS |volume=101 |issue=21 |pages=8066–71 |date=May 2004 |pmid=15148395 |pmc=419558 |doi=10.1073/pnas.0308602101 |doi-access=free }}</ref> Except for the ] and three species in the genus '']'' in the phylum ], they are all non-photosynthethic, but many foraminifera and radiolaria have a symbiotic relationship with unicellular algae.<ref>{{Cite journal |last1=Gast |first1=Rebecca J.|last2=Caron |first2=David A. |date=2001-10-01 |title=Photosymbiotic associations in planktonic foraminifera and radiolaria |journal=Hydrobiologia |volume=461 |issue=1 |pages=1–7 |doi=10.1023/A:1012710909023 |s2cid=1387879}}</ref> A multicellular form, ''Guttulinopsis vulgaris'', a cellular ], has been described.<ref>{{cite journal |last=Brown |first=Matthew W. |last2=Kolisko |first2=Martin |last3=Silberman |first3=Jeffrey D. |last4=Roger |first4=Andrew J. |title=Aggregative Multicellularity Evolved Independently in the Eukaryotic Supergroup Rhizaria |journal=] |volume=22 |issue=12 |year=2012 |doi=10.1016/j.cub.2012.04.021 |pages=1123–1127|doi-access=free }}</ref> This group was used by ] in 2002, although the term "Rhizaria" had been long used for clades within the currently recognized taxon. Being described mainly from ] sequences, they vary considerably in form, having no clear morphological distinctive characters (]), but for the most part they are ]s with ], ], or ]-supported ]s. In the absence of an apomorphy, the group is ill-defined, and its composition has been very fluid. Some Rhizaria possess mineral exoskeletons (] or ]s), which are in different clades within Rhizaria made out of ] ({{chem2|SiO2}}), ] ({{chem2|SrSO4}}), or ] ({{chem2|CaCO3}}). Certain species 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.<ref>Caron, D. (2016). The rise of Rhizaria. Nature (London), 532(7600), 444–445. https://doi.org/10.1038/nature17892</ref> |
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This group was used by ] in 2002, although the term "Rhizaria" had been long used for clades within the currently recognized taxon. Being described mainly from ] sequences, they vary considerably in form, having no clear morphological distinctive characters (]), but for the most part they are ]s with ], ], or ]-supported ]s. In the absence of an apomorphy, the group is ill-defined, and its composition has been very fluid. Some Rhizaria possess mineral exoskeletons (] or ]s), which are in different clades within Rhizaria made out of ] ({{chem2|SiO2}}), ] ({{chem2|SrSO4}}), or ] ({{chem2|CaCO3}}). Certain species 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.<ref>Caron, D. (2016). The rise of Rhizaria. Nature (London), 532(7600), 444–445. https://doi.org/10.1038/nature17892</ref> |
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==Groups== |
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==Groups== |
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A few other groups may be included in the Cercozoa, but some trees appear closer to the Foraminifera. These are the ] and ], parasites of plants and animals, respectively, and the peculiar amoeba '']''. 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 ] in 2002,<ref>{{cite journal | first=Thomas | last=Cavalier-Smith | author-link=Thomas Cavalier-Smith | title=The phagotrophic origin of eukaryotes and phylogenetic classification of Protozoa | journal=International Journal of Systematic and Evolutionary Microbiology | year=2002 |volume=52 |issue=2 |pages=297–354 | issn=1466-5026 | url=http://ijs.sgmjournals.org/cgi/content/abstract/52/2/297 | access-date=2007-06-08 | pmid=11931142 | doi=10.1099/00207713-52-2-297}}</ref> who also included the ]s and ]. |
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A few other groups may be included in the Cercozoa, but some trees appear closer to the Foraminifera. These are the ] and ], parasites of plants and animals, respectively, and the peculiar amoeba '']''. 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 ] in 2002,<ref>{{cite journal | first=Thomas | last=Cavalier-Smith | author-link=Thomas Cavalier-Smith | title=The phagotrophic origin of eukaryotes and phylogenetic classification of Protozoa | journal=International Journal of Systematic and Evolutionary Microbiology | year=2002 |volume=52 |issue=2 |pages=297–354 | issn=1466-5026 | url=http://ijs.sgmjournals.org/cgi/content/abstract/52/2/297 | access-date=2007-06-08 | pmid=11931142 | doi=10.1099/00207713-52-2-297}}</ref> who also included the ]s and ]. |
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A noteworthy order that belongs to ] is the ].<ref name=Hartikainen2014>{{cite journal |last1=Hartikainen |first1=H. |last2=Stentiford |first2=G.D. |last3=Bateman |first3=K.S. |last4=Berney |first4=C. |last5=Feist |first5=S.W. |last6=Longshaw |first6=M. |last7=Okamura |first7=B. |last8=Stone |first8=D. |last9=Ward |first9=G. |last10=Wood |first10=C. |last11=Bass |first11=D. |year=2014 |title=Mikrocytids are a broadly distributed and divergent radiation of parasites in aquatic invertebrates |url =https://nhm.openrepository.com/bitstream/10141/622246/1/Mikrocytids_CurrBiol_2014.pdf |journal=Curr Biol |volume=24 |issue=7| pages=807–12 |doi=10.1016/j.cub.2014.02.033 |pmid=24656829| s2cid=17180719 |doi-access=free }}</ref> These are parasites of ]s. This includes the causative agent of Denman Island Disease, ''Mikrocytos mackini'' a small (2−3 μm diameter) amitochondriate protistan.<ref>{{Cite journal |last1=Hine |first1=P.M. |last2=Bower |first2=S.M. |last3=Meyer |first3=G.R. |last4=Cochennec-Laureau |first4=N. |last5=Berthe |first5=F.C.J. |date=2001 |title=Ultrastructure of Mikrocytos mackini, the cause of Denman Island disease in oysters Crassostrea spp. and Ostrea spp. in British Columbia, Canada |url=http://www.int-res.com/abstracts/dao/v45/n3/p215-227/ |journal=Diseases of Aquatic Organisms |volume=45 |issue=3 |pages=215–227 |doi=10.3354/dao045215 |pmid=11558731 |issn=0177-5103 |doi-access=free}}</ref> |
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A noteworthy order that belongs to ] is the ].<ref name=Hartikainen2014>{{cite journal |last1=Hartikainen |first1=H. |last2=Stentiford |first2=G.D. |last3=Bateman |first3=K.S. |last4=Berney |first4=C. |last5=Feist |first5=S.W. |last6=Longshaw |first6=M. |last7=Okamura |first7=B. |last8=Stone |first8=D. |last9=Ward |first9=G. |last10=Wood |first10=C. |last11=Bass |first11=D. |year=2014 |title=Mikrocytids are a broadly distributed and divergent radiation of parasites in aquatic invertebrates |url =https://nhm.openrepository.com/bitstream/10141/622246/1/Mikrocytids_CurrBiol_2014.pdf |journal=Curr Biol |volume=24 |issue=7| pages=807–12 |doi=10.1016/j.cub.2014.02.033 |pmid=24656829| s2cid=17180719 |doi-access=free }}</ref> These are parasites of ]s. This includes the causative agent of Denman Island Disease, ''Mikrocytos mackini'' a small (2−3 μm diameter) amitochondriate protistan.<ref>{{Cite journal |last1=Hine |first1=P.M. |last2=Bower |first2=S.M. |last3=Meyer |first3=G.R. |last4=Cochennec-Laureau |first4=N. |last5=Berthe |first5=F.C.J. |date=2001 |title=Ultrastructure of Mikrocytos mackini, the cause of Denman Island disease in oysters Crassostrea spp. and Ostrea spp. in British Columbia, Canada |url=http://www.int-res.com/abstracts/dao/v45/n3/p215-227/ |journal=Diseases of Aquatic Organisms |volume=45 |issue=3 |pages=215–227 |doi=10.3354/dao045215 |pmid=11558731 |issn=0177-5103 |doi-access=free}}</ref> |
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==Evolutionary relationships== |
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==Evolutionary relationships== |
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Rhizaria are part of the ] (Stramenopiles, Alveolates, Rhizaria), a grouping that had been presaged in 1993 through a study of mitochondrial morphologies.<ref>Seravin LN. Osnovnye tipy i formy tonkogo stroeniia krist mitokhondriĭ: stepen' ikh évoliutsionnoĭ stabil'nosti (sposobnost' k morfologicheskim transformatsiiam) . Tsitologiia. 1993;35(4):3-34. Russian. PMID 8328023.</ref> SAR is currently placed in the ] along with ], ], ], and several minor clades. |
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Rhizaria are part of the ] (Stramenopiles, Alveolates, Rhizaria), a grouping that had been presaged in 1993 through a study of mitochondrial morphologies.<ref>Seravin LN. Osnovnye tipy i formy tonkogo stroeniia krist mitokhondriĭ: stepen' ikh évoliutsionnoĭ stabil'nosti (sposobnost' k morfologicheskim transformatsiiam) . Tsitologiia. 1993;35(4):3-34. Russian. PMID 8328023.</ref> SAR is currently placed in the ] along with ], ], ], and several minor clades. |
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Historically, many rhizarians were considered ]s because of their motility and ]. However, when a simple animal-plant dichotomy was superseded by a recognition of additional kingdoms, taxonomists generally placed amoebae in the kingdom ]. When scientists began examining the evolutionary relationships among eukaryotes in the 1970s, it became clear that the kingdom ] was ]. Rhizaria appear to share a common ancestor with ] and ] forming part of the SAR (Stramenopiles+Alveolates+Rhizaria) super assemblage.<ref>{{cite journal |title=Phylogenomics Reshuffles the Eukaryotic Supergroups| journal=]| year=2007 |volume=2 |pages=e790– |issue=8| doi=10.1371/journal.pone.0000790 |pmid=17726520 |last1=Burki |first1=F. |last2=Shalchian-Tabrizi |first2=K. |last3=Minge |first3=M. |last4=Skjaeveland |first4=A. |last5=Nikolaev |first5=S.I. |last6=Jakobsen |first6=K.S. |last7=Pawlowski |first7=J. |pmc=1949142 |editor1-last=Butler |editor1-first=Geraldine |
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Historically, many rhizarians were considered ]s because of their motility and ]. However, when a simple animal-plant dichotomy was superseded by a recognition of additional kingdoms, taxonomists generally placed amoebae in the kingdom ]. When scientists began examining the evolutionary relationships among eukaryotes in the 1970s, it became clear that the kingdom ] was ]. Rhizaria appear to share a common ancestor with ] and ] forming part of the SAR super assemblage.<ref>{{cite journal |title=Phylogenomics Reshuffles the Eukaryotic Supergroups| journal=]| year=2007 |volume=2 |pages=e790– |issue=8| doi=10.1371/journal.pone.0000790 |pmid=17726520 |last1=Burki |first1=F. |last2=Shalchian-Tabrizi |first2=K. |last3=Minge |first3=M. |last4=Skjaeveland |first4=A. |last5=Nikolaev |first5=S.I. |last6=Jakobsen |first6=K.S. |last7=Pawlowski |first7=J. |pmc=1949142 |editor1-last=Butler |editor1-first=Geraldine |
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| bibcode=2007PLoSO...2..790B| doi-access=free}}</ref> Rhizaria has been supported by molecular phylogenetic studies as a monophyletic group.<ref name=Burki2008>{{cite journal |last1=Burki |first1=Fabien |last2=Shalchian-Tabrizi |first2=Kamran |last3=Pawlowski |first3=Jan |title=Phylogenomics reveals a new 'megagroup' including most photosynthetic eukaryotes |journal=Biology Letters |date=August 23, 2008 |doi=10.1098/rsbl.2008.0224 |pmid=18522922 |volume=4 |issue=4 |pmc=2610160 |pages=366–369}}</ref> Biosynthesis of ] precursors in various rhizaria<ref>{{Cite journal |last1=Hallmann |first1=Christian |last2=Stuhr |first2=Marleen |last3=Kucera |first3=Michal |last4=Zonneveld |first4=Karin |last5=Bobrovskiy |first5=Ilya |last6=Bowser |first6=Samuel S. |last7=Pawlowski |first7=Jan |last8=Deckker |first8=Patrick D |last9=Nowack |first9=Eva C. M. |display-authors=3 |date=2019-03-04 |title=Putative sponge biomarkers in unicellular Rhizaria question an early rise of animals |journal=Nature Ecology & Evolution |volume=3 |issue=4 |pages=577–581 |doi=10.1038/s41559-019-0806-5 |pmid=30833757 |s2cid=71148672}}</ref> suggests a relevant ecological role already during the ]. |
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| bibcode=2007PLoSO...2..790B| doi-access=free}}</ref> Rhizaria has been supported by molecular phylogenetic studies as a monophyletic group.<ref name=Burki2008>{{cite journal |last1=Burki |first1=Fabien |last2=Shalchian-Tabrizi |first2=Kamran |last3=Pawlowski |first3=Jan |title=Phylogenomics reveals a new 'megagroup' including most photosynthetic eukaryotes |journal=Biology Letters |date=August 23, 2008 |doi=10.1098/rsbl.2008.0224 |pmid=18522922 |volume=4 |issue=4 |pmc=2610160 |pages=366–369}}</ref> Biosynthesis of ] precursors in various rhizaria<ref>{{Cite journal |last1=Hallmann |first1=Christian |last2=Stuhr |first2=Marleen |last3=Kucera |first3=Michal |last4=Zonneveld |first4=Karin |last5=Bobrovskiy |first5=Ilya |last6=Bowser |first6=Samuel S. |last7=Pawlowski |first7=Jan |last8=Deckker |first8=Patrick D |last9=Nowack |first9=Eva C. M. |display-authors=3 |date=2019-03-04 |title=Putative sponge biomarkers in unicellular Rhizaria question an early rise of animals |journal=Nature Ecology & Evolution |volume=3 |issue=4 |pages=577–581 |doi=10.1038/s41559-019-0806-5 |pmid=30833757 |s2cid=71148672}}</ref> suggests a relevant ecological role already during the ]. |
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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, who also included the centrohelids and Apusozoa.
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