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{{Short description|Infrakingdom of protists}} {{Short description|Infrakingdom of protists}}
{{Automatic taxobox {{Automatic taxobox
| fossil_range = {{long fossil range|Neoproterozoic|recent|] - ]}} | fossil_range={{long fossil range|650|0|650 Mya<ref name="RhizariaPhylo2018"/> (]) - ]}}
| image = Ammonia tepida.jpg | image=Ammonia tepida.jpg
| image_caption = '']'' (]) | image_caption='']'' (])
| taxon = Rhizaria | taxon=Rhizaria
| authority = ], 2002 | authority=], 2002
| subdivision_ranks = Phyla | subdivision_ranks=Phyla
|subdivision_ref = <ref name="Adl 2019">{{cite Q|Q57086550}}</ref>
| subdivision = | subdivision =
*] * ]
**] * ]
**] * ]
⚫ | display_parents=6
*]
**]
**]
⚫ | display_parents = 6
}} }}


The '''Rhizaria''' are an ill-defined but species-rich supergroup of mostly ]<ref>{{cite web | url = http://www.palaeos.com/Eukarya/Units/Rhizaria/Rhizaria.html | year = 2004 | author = Christopher Taylor | 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 |vauthors=Nikolaev SI, Berney C, Fahrni JF, etal |title=The twilight of Heliozoa and rise of Rhizaria, an emerging supergroup of amoeboid eukaryotes |journal=Proc. Natl. Acad. Sci. U.S.A. |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|language=en|volume=461|issue=1|pages=1–7|doi=10.1023/A:1012710909023|s2cid=1387879|issn=1573-5117}}</ref> A multicellular form, ''Guttulinopsis vulgaris'', a cellular ], has also been described.<ref>{{cite journal | year = 2012 | author = Brown| title = Aggregative Multicellularity Evolved Independently in the Eukaryotic Supergroup Rhizaria |display-authors=etal | doi=10.1016/j.cub.2012.04.021 | volume=22 | issue = 12 | journal=Current Biology | pages=1123–1127 | pmid=22608512| doi-access=free }}</ref> The '''Rhizaria''' are a diverse and species-rich ] 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 |last1=Nikolaev |first1=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-photosynthetic, but many ] and ] 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 |last1=Brown |first1=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 |doi=10.1016/j.cub.2012.04.021 |pages=1123–7 |date=June 2012 |pmid=22608512 |doi-access=free |bibcode=2012CBio...22.1123B }}</ref> This group was used by ] in 2002, although the term "Rhizaria" had been long used for clades within the currently recognized taxon.

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&nbsp;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> 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&nbsp;cm in diameter and greater than 1&nbsp;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>{{cite journal |vauthors=Caron DA |title=Ocean science: The rise of Rhizaria |journal=Nature |volume=532 |issue=7600 |pages=444–5 |date=April 2016 |pmid=27096370 |doi=10.1038/nature17892 |bibcode=2016Natur.532..444C }}</ref>


==Groups== ==Groups==
{{Further|wikispecies:Rhizaria}} {{Further|wikispecies:Rhizaria}}

The three main groups of Rhizaria are:<ref name="pmid17174576">{{cite journal |vauthors=Moreira D, von der Heyden S, Bass D, López-García P, Chao E, Cavalier-Smith T |title=Global eukaryote phylogeny: Combined small- and large-subunit ribosomal DNA trees support monophyly of Rhizaria, Retaria and Excavata |journal=Mol. Phylogenet. Evol. |volume=44 |issue=1 |pages=255–66 |date=July 2007 |pmid=17174576 |doi=10.1016/j.ympev.2006.11.001 }}</ref> The three main groups of Rhizaria are:<ref name="pmid17174576">{{cite journal |vauthors=Moreira D, von der Heyden S, Bass D, López-García P, Chao E, Cavalier-Smith T |title=Global eukaryote phylogeny: Combined small- and large-subunit ribosomal DNA trees support monophyly of Rhizaria, Retaria and Excavata |journal=Mol. Phylogenet. Evol. |volume=44 |issue=1 |pages=255–66 |date=July 2007 |pmid=17174576 |doi=10.1016/j.ympev.2006.11.001 |bibcode=2007MolPE..44..255M }}</ref>


* ] – various amoebae and flagellates, usually with ] and common in soil * ] – various amoebae and flagellates, usually with ] and common in soil
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* ] – amoeboids with ]s, common as marine ] * ] – amoeboids with ]s, common as marine ]


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 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 | url=http://ijs.sgmjournals.org/cgi/content/abstract/52/2/297 | pmid=11931142 | doi=10.1099/00207713-52-2-297}}</ref> who also included the ]s and ].
| 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 ].


A noteworthy order that belongs to ] is the ].<ref name=Hartikainen2014>{{cite journal | last1 = Hartikainen | first1 = H | last2 = Stentiford | first2 = GD | last3 = Bateman | first3 = KS | last4 = Berney | first4 = C | last5 = Feist | first5 = SW | 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=Pm|last2=Bower|first2=Sm|last3=Meyer|first3=Gr|last4=Cochennec-Laureau|first4=N|last5=Berthe|first5=Fcj|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|language=en|volume=45|issue=3|pages=215–227|doi=10.3354/dao045215|pmid=11558731|issn=0177-5103|doi-access=free}}</ref> 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 |bibcode=2014CBio...24..807H }}</ref> These are parasites of ]s. This includes the causative agent of Denman Island Disease, ''Mikrocytos mackini'' a small (2−3&nbsp;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 |doi-access=free}}</ref>

== History ==

Similarities between various Rhizaria organisms have been noticed since the 19th century. In his 1861 classification of the ] (amoebae), the zoologist ] proposed the order '''Reticularia''', which consisted of ] and ] on the basis of their very similar thin, reticulose pseudopodia with granules circulating inside.<ref name="Carpenter 1861">{{Cite journal|last=Carpenter|first=William Benjamin|date=1861|title=XLVII.-On the systematic arrangement of the Rhizopoda|url=https://www.biodiversitylibrary.org/item/30158#page/478/mode/1up|journal=Natural History Review (Dublin and London)|volume=1|issue=4|pages=478}}</ref> However, the idea that these organisms and others such as ] were all related to one another emerged rather recently, with the help of molecular phylogenetics and advanced microscopy techniques in the late 20th century.<ref name="Burki 2014">{{cite journal|vauthors=Burki F, Keeling PJ|date=February 2014|title=Rhizaria|journal=Current Biology|volume=24|issue=3|pages=R103–7 |pmid=24502779 |doi=10.1016/j.cub.2013.12.025|doi-access=free|bibcode=2014CBio...24.R103B }}</ref>


==Evolutionary relationships== ==Evolutionary relationships==
{{See also|Eukaryote#Phylogeny}} {{See also|Eukaryote#Phylogeny}}
⚫ 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.


⚫ Rhizaria are part of the ] (Stramenopiles, Alveolates, Rhizaria), a grouping that had been presaged in 1993 through a study of mitochondrial morphologies.<ref>{{cite journal |vauthors=Seravin LN |title= |language=Russian |journal=Tsitologiia |volume=35 |issue=4 |pages=3–34 |date=1993 |pmid=8328023 }}</ref> SAR is currently placed in the ] along with ], ], ], and several minor clades.
⚫ 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 1970's, 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 = SI | last6 = Jakobsen | first6 = KS | last7 = Pawlowski | first7 = J | pmc = 1949142 | editor1-last = Butler | editor1-first = Geraldine
⚫ | 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=366–9}}</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 De|last9=Nowack|first9=Eva C. M.|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|language=en|pages=577–581|doi=10.1038/s41559-019-0806-5|pmid=30833757|s2cid=71148672|issn=2397-334X}}</ref> suggests a relevant ecological role already during the ].


⚫ 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
⚫ ===Phylogeny===
⚫ | 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=366–9}}</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 |bibcode=2019NatEE...3..577N |s2cid=71148672}}</ref> suggests a relevant ecological role already during the ].


⚫ ===Phylogeny===
⚫ Rhizaria is a ] composed of two sister phyla: ] and ]. Subsequently, Cercozoa and Retaria are also ].<ref name="pmid18952499">{{cite journal |vauthors=Bass D, Chao EE, Nikolaev S |title=Phylogeny of Novel Naked Filose and Reticulose Cercozoa: Granofilosea cl. n. and Proteomyxidea Revised |journal=Protist |volume=160 |issue=1 |pages=75–109 |date=February 2009 |pmid=18952499 |doi=10.1016/j.protis.2008.07.002 |display-authors=etal}}</ref><ref name=Howe>{{citation | date=2011| author = Howe | title = Novel Cultured Protists Identify Deep-branching Environmental DNA Clades of Cercozoa: New Genera Tremula, Micrometopion, Minimassisteria, Nudifila, Peregrinia| volume=162 |issue= 2|pages=332–372 |journal=Protist | doi=10.1016/j.protis.2010.10.002 |display-authors=etal | pmid=21295519}}</ref> The following cladogram depicts the evolutionary relationships between all rhizarian ]es, and is made after the works of ] ''et al.'' (2018),<ref name="RhizariaPhylo2018">{{cite journal |title=Multigene phylogeny and cell evolution of chromist infrakingdom Rhizaria: contrasting cell organisation of sister phyla Cercozoa and Retaria |first1=Thomas |last1=Cavalier-Smith |author1-link=Thomas Cavalier-Smith |first2=Ema E .|last2=Chao |first3=Rhodri |last3=Lewis |date=April 2018 |journal=Protoplasma |volume=255 |issue=5 |pages=1517–1574|doi=10.1007/s00709-018-1241-1 |pmid=29666938 |pmc=6133090}}</ref> Irwin ''et al.'' (2019)<ref name="MonoCercozoa">{{Cite journal |date=2019-01-01 |title=Phylogenomics supports the monophyly of the Cercozoa |journal=Molecular Phylogenetics and Evolution |volume=130 |pages=416–423 |doi=10.1016/j.ympev.2018.09.004 |pmid=30318266 |issn=1055-7903 |last1=Irwin |first1=Nicholas A. T. |last2=Tikhonenkov|first2=Denis V.|last3=Hehenberger |first3=Elisabeth |last4=Mylnikov |first4=Alexander P. |last5=Burki |first5=Fabien |last6=Keeling |first6=Patrick J. |display-authors=3 |s2cid=52982396}}</ref> and Sierra ''et al.'' (2022):<ref name="TaxonRichForams">{{cite journal |last=Sierra |first=R |display-authors=etal |title=Taxon-rich transcriptomics supports higher-level phylogeny and major evolutionary trends in Foraminifera |journal=Molecular Phylogenetics and Evolution |volume=174 |date=2022 |pages=107546 |issn=1055-7903 |doi=10.1016/j.ympev.2022.107546 |pmid=35690380 |doi-access=free}}</ref><!--and if they don't agree? this isn't ok-->


⚫ Rhizaria is a ] composed of two sister phyla: ] and ]. Subsequently, Cercozoa and Retaria are also ].<ref name="pmid18952499">{{cite journal |last1=Bass |first1=D. |last2=Chao |first2=E.E. |last3=Nikolaev |first3=S. |title=Phylogeny of Novel Naked Filose and Reticulose Cercozoa: Granofilosea cl. n. and Proteomyxidea Revised |journal=Protist |volume=160 |issue=1 |pages=75–109 |date=February 2009 |pmid=18952499 |doi=10.1016/j.protis.2008.07.002 |display-authors=etal}}</ref><ref name=Howe>>{{cite journal | last1=Howe | first1=Alexis T. | last2=Bass | first2=David | last3=Scoble | first3=Josephine M. | last4=Lewis | first4=Rhodri | last5=Vickerman | first5=Keith | last6=Arndt | first6=Hartmut | last7=Cavalier-Smith | first7=Thomas |author7-link=Thomas Cavalier-Smith |display-authors=3 |title=Novel Cultured Protists Identify Deep-branching Environmental DNA Clades of Cercozoa: New Genera Tremula, Micrometopion, Minimassisteria, Nudifila, Peregrinia |date=2011 |volume=162 |issue=2 |pages=332–372 |journal=Protist |doi=10.1016/j.protis.2010.10.002 |pmid=21295519}}</ref> The following cladogram depicts the evolutionary relationships between all rhizarian ]es, and is made after the works of ] ''et al.'' (2018),<ref name="RhizariaPhylo2018">{{cite journal |title=Multigene phylogeny and cell evolution of chromist infrakingdom Rhizaria: contrasting cell organisation of sister phyla Cercozoa and Retaria |first1=Thomas |last1=Cavalier-Smith |author1-link=Thomas Cavalier-Smith |first2=Ema E . |last2=Chao |first3=Rhodri |last3=Lewis |date=April 2018 |journal=Protoplasma |volume=255 |issue=5 |pages=1517–1574|doi=10.1007/s00709-018-1241-1 |pmid=29666938 |pmc=6133090}}</ref> and Irwin ''et al.'' (2019).<ref name="MonoCercozoa">{{Cite journal |date=2019-01-01 |title=Phylogenomics supports the monophyly of the Cercozoa |journal=Molecular Phylogenetics and Evolution |volume=130 |pages=416–423 |doi=10.1016/j.ympev.2018.09.004 |pmid=30318266 |last1=Irwin |first1=Nicholas A. T. |last2=Tikhonenkov|first2=Denis V.|last3=Hehenberger |first3=Elisabeth |last4=Mylnikov |first4=Alexander P. |last5=Burki |first5=Fabien |last6=Keeling |first6=Patrick J. |bibcode=2019MolPE.130..416I |display-authors=3 |s2cid=52982396}}</ref>


{{clade {{clade
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|2=] ] |2=] (paraphyletic) ]
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==Sexual cycle==

Complete sexual life cycles have been demonstrated for two lineages (] and '']'') and direct evidence for ] or ] has been observed in five lineages ('']'', '']'', ], ] and '']'').<ref name="Lahr2011">{{cite journal |vauthors=Lahr DJ, Parfrey LW, Mitchell EA, Katz LA, Lara E |title=The chastity of amoebae: re-evaluating evidence for sex in amoeboid organisms |journal=Proc Biol Sci |volume=278 |issue=1715 |pages=2081–90 |date=July 2011 |pmid=21429931 |pmc=3107637 |doi=10.1098/rspb.2011.0289 }}</ref> In particular, the Foraminifera are marine ]e that are defined by a dynamic network of ], and the production of intricate shells.<ref name =Lahr2011/> These amoeba have complex sexual life cycles with meiosis and ] production occurring at separate stages.<ref name =Lahr2011/>


== References == == References ==
{{Reflist|2}} {{Reflist|30em}}


== External links == == External links ==

* {{dead link|date=December 2024|bot=medic}}{{cbignore|bot=medic}} at UniEuk Taxonomy Map
* *
* *
* https://www-nature-com.


{{Eukaryota}} {{Eukaryota}}

Latest revision as of 05:46, 21 December 2024

Infrakingdom of protists

Rhizaria
Temporal range: 650 Mya (Neoproterozoic) - Present Pha. Proterozoic Archean Had.
Ammonia tepida (Foraminifera)
Scientific classification Edit this classification
Domain: Eukaryota
Clade: Diaphoretickes
Clade: TSAR
Clade: SAR
Clade: Rhizaria
Cavalier-Smith, 2002
Phyla

The Rhizaria are a diverse and species-rich supergroup of mostly unicellular eukaryotes. Except for the Chlorarachniophytes and three species in the genus Paulinella in the phylum Cercozoa, they are all non-photosynthetic, but many Foraminifera and Radiolaria have a symbiotic relationship with unicellular algae. A multicellular form, Guttulinopsis vulgaris, a cellular slime mold, has been described. 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 exoskeletons (thecae or loricas), which are in different clades within Rhizaria made out of opal (SiO2), celestite (SrSO4), or calcite (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.

Groups

Further information: wikispecies:Rhizaria

The three main groups of Rhizaria are:

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.

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

History

Similarities between various Rhizaria organisms have been noticed since the 19th century. In his 1861 classification of the Rhizopoda (amoebae), the zoologist William B. Carpenter proposed the order Reticularia, which consisted of Foraminifera and Gromiida on the basis of their very similar thin, reticulose pseudopodia with granules circulating inside. However, the idea that these organisms and others such as Radiolaria were all related to one another emerged rather recently, with the help of molecular phylogenetics and advanced microscopy techniques in the late 20th century.

Evolutionary relationships

See 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. 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 1970s, 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 super assemblage. Rhizaria has been supported by molecular phylogenetic studies as a monophyletic group. Biosynthesis of 24-isopropyl cholestane precursors in various rhizaria 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. The following cladogram depicts the evolutionary relationships between all rhizarian classes, and is made after the works of Cavalier-Smith et al. (2018), and Irwin et al. (2019).

SAR Supergroup
Rhizaria
Cercozoa
Filosa
Monadofilosa

Imbricatea

Thecofilosea

Sarcomonadea (paraphyletic)

Helkesea

Metromonadea

Reticulofilosa

Granofilosea

Chlorarachnea

Endomyxa

Phytomyxea

Gromiidea

Lapot gusevi

Retaria

Foraminifera

Polycystinea

Acantharea

Sticholonchea

Halvaria

Stramenopiles

Alveolata

Sexual cycle

Complete sexual life cycles have been demonstrated for two lineages (Foraminifera and Gromia) and direct evidence for karyogamy or meiosis has been observed in five lineages (Euglyphida, Thecofilosea, Chlorarachniophyta, Plasmodiophorida and Phaeodarea). In particular, the Foraminifera are marine amoebae that are defined by a dynamic network of pseudopodia, and the production of intricate shells. These amoeba have complex sexual life cycles with meiosis and gamete production occurring at separate stages.

References

  1. ^ 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.
  2. Sina M. Adl; David Bass; Christopher E. Lane; et al. (1 January 2019). "Revisions to the Classification, Nomenclature, and Diversity of Eukaryotes". Journal of Eukaryotic Microbiology. 66 (1): 4–119. doi:10.1111/JEU.12691. ISSN 1066-5234. PMC 6492006. PMID 30257078. Wikidata Q57086550.
  3. Taylor, Christopher (2004). "Rhizaria". Archived from the original on 2009-04-20.
  4. Nikolaev, Sergey I.; Berney, Cédric; Fahrni, José F.; et al. (May 2004). "The twilight of Heliozoa and rise of Rhizaria, an emerging supergroup of amoeboid eukaryotes". PNAS. 101 (21): 8066–71. doi:10.1073/pnas.0308602101. PMC 419558. PMID 15148395.
  5. 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. S2CID 1387879.
  6. Brown, Matthew W.; Kolisko, Martin; Silberman, Jeffrey D.; Roger, Andrew J. (June 2012). "Aggregative Multicellularity Evolved Independently in the Eukaryotic Supergroup Rhizaria". Current Biology. 22 (12): 1123–7. Bibcode:2012CBio...22.1123B. doi:10.1016/j.cub.2012.04.021. PMID 22608512.
  7. Caron DA (April 2016). "Ocean science: The rise of Rhizaria". Nature. 532 (7600): 444–5. Bibcode:2016Natur.532..444C. doi:10.1038/nature17892. PMID 27096370.
  8. 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. Bibcode:2007MolPE..44..255M. doi:10.1016/j.ympev.2006.11.001. PMID 17174576.
  9. 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. PMID 11931142.
  10. Hartikainen, H.; Stentiford, G.D.; Bateman, K.S.; Berney, C.; Feist, S.W.; 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. Bibcode:2014CBio...24..807H. doi:10.1016/j.cub.2014.02.033. PMID 24656829. S2CID 17180719.
  11. Hine, P.M.; Bower, S.M.; Meyer, G.R.; Cochennec-Laureau, N.; Berthe, F.C.J. (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. PMID 11558731.
  12. Carpenter, William Benjamin (1861). "XLVII.-On the systematic arrangement of the Rhizopoda". Natural History Review (Dublin and London). 1 (4): 478.
  13. Burki F, Keeling PJ (February 2014). "Rhizaria". Current Biology. 24 (3): R103–7. Bibcode:2014CBio...24.R103B. doi:10.1016/j.cub.2013.12.025. PMID 24502779.
  14. Seravin LN (1993). "". Tsitologiia (in Russian). 35 (4): 3–34. PMID 8328023.
  15. Burki, F.; Shalchian-Tabrizi, K.; Minge, M.; Skjaeveland, A.; Nikolaev, S.I.; Jakobsen, K.S.; 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.
  16. 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.
  17. Hallmann, Christian; Stuhr, Marleen; Kucera, Michal; et al. (2019-03-04). "Putative sponge biomarkers in unicellular Rhizaria question an early rise of animals". Nature Ecology & Evolution. 3 (4): 577–581. Bibcode:2019NatEE...3..577N. doi:10.1038/s41559-019-0806-5. PMID 30833757. S2CID 71148672.
  18. Bass, D.; Chao, E.E.; 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.
  19. >Howe, Alexis T.; Bass, David; Scoble, Josephine M.; 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.
  20. Irwin, Nicholas A. T.; Tikhonenkov, Denis V.; Hehenberger, Elisabeth; et al. (2019-01-01). "Phylogenomics supports the monophyly of the Cercozoa". Molecular Phylogenetics and Evolution. 130: 416–423. Bibcode:2019MolPE.130..416I. doi:10.1016/j.ympev.2018.09.004. PMID 30318266. S2CID 52982396.
  21. ^ Lahr DJ, Parfrey LW, Mitchell EA, Katz LA, Lara E (July 2011). "The chastity of amoebae: re-evaluating evidence for sex in amoeboid organisms". Proc Biol Sci. 278 (1715): 2081–90. doi:10.1098/rspb.2011.0289. PMC 3107637. PMID 21429931.

External links

Eukaryote classification
Domain
Archaea
Bacteria
Eukaryota
(major groups
Excavata
Diaphoretickes
Hacrobia
Cryptista
Rhizaria
Alveolata
Stramenopiles
Plants
Amorphea
Amoebozoa
Opisthokonta
Animals
Fungi
Mesomycetozoea)
Amorphea
Amoebozoa
   Obazoa   
Opisthokonta
Holomycota
   Holozoa   
Filozoa
Choanozoa
Diaphoretickes
Haptista
    TSAR    
     SAR     
Rhizaria
Alveolata
Myzozoa
Stramenopiles
Bigyra
Gyrista
     CAM     
Pancryptista
Cryptista
Archaeplastida
Viridiplantae
(Plants sensu lato)
Streptophyta
Excavates*
Discoba
Discicristata
Metamonada
Malawimonada
Incertae sedis
Hemimastigophora
Ancyromonadida
CRuMs
Rhizaria classification
Domain
Archaea
Bacteria
Eukaryota
(major groups
Excavata
Diaphoretickes
Hacrobia
Cryptista
Rhizaria
Alveolata
Stramenopiles
Plants
Amorphea
Amoebozoa
Opisthokonta
Animals
Fungi
Mesomycetozoea)
 Cercozoa 
Reticulofilosa*
Skiomonadea
Granofilosea
Chlorarachnea
Monadofilosa
Eoglissa
Metromonadea
Helkesea
Ventrifilosa
Sarcomonadea*
Imbricatea
Silicofilosea
Thecofilosea
Endomyxa
Marimyxia
Gromiidea
Ascetosporea
Proteomyxia
   Retaria   
Radiolaria
Sticholonchea
Acantharea
Polycystinea
Foraminifera
Monothalamea*
Tubothalamea
Globothalamea
Incertae sedis
Incertae sedis
Taxon identifiers
Rhizaria
Categories: