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The '''Rhizaria''' are a 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 | archiveurl = https://web.archive.org/web/20090420034036/http://www.palaeos.com/Eukarya/Units/ but the averageis500years
The '''Rhizaria''' are a 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 | archiveurl = https://web.archive.org/web/20090420034036/http://www.palaeos.com/Eukarya/Units/Rhizaria/Rhizaria.html | archivedate = 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 }}</ref> Except from the ] and three species in the genus ] in the phylum ], they are all non-photosyntethic, but many foraminifers have a symbiotic relationship with unicellular algae. A multicellular form, ''Guttulinopsis vulgaris'', a cellular ], has also been described.<ref>{{cite journal | url=http://www.cell.com/current-biology/abstract/S0960-9822(12)00413-7 | 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}}</ref>
Rhizaria/Rhizaria.html | archivedate = 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 }}</ref> Except from the ] and three species in the genus ] in the phylum ], they are all non-photosyntethic, but many foraminifers have a symbiotic relationship with unicellular algae. A multicellular form, ''Guttulinopsis vulgaris'', a cellular ], has also been described.<ref>{{cite journal | url=http://www.cell.com/current-biology/abstract/S0960-9822(12)00413-7 | 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}}</ref>
This supergroup was proposed by ] in 2002. 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 filose, reticulose, or microtubule-supported ]s. Many produce shells or skeletons, which may be quite complex in structure, and these make up the vast majority of protozoan fossils. Nearly all have ] with tubular ]e.
This supergroup was proposed by ] in 2002. 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 filose, reticulose, or microtubule-supported ]s. Many produce shells or skeletons, which may be quite complex in structure, and these make up the vast majority of protozoan fossils. Nearly all have ] with tubular ]e.
The Rhizaria are a species-rich supergroup of mostly unicellulareukaryotes. Except from the Chlorarachniophyte and three species in the genus Paulinella in the phylum Cercozoa, they are all non-photosyntethic, but many foraminifers have a symbiotic relationship with unicellular algae. A multicellular form, Guttulinopsis vulgaris, a cellular slime mold, has also been described.
This supergroup was proposed by Cavalier-Smith in 2002. 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. Many produce shells or skeletons, which may be quite complex in structure, and these make up the vast majority of protozoan fossils. Nearly all have mitochondria with tubular cristae.
A few other groups may be included in the Cercozoa, but on 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.
Another order that appears to belong to this taxon is the Mikrocytida. These are parasites of oysters.
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 rhizarians in the kingdom Protista. When scientists began examining the evolutionary relationships among eukaryotes using molecular data, 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. 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
Phylogeny based on Bass et al. 2009, Howe et al. 2011, and Silar 2016.
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. PMID17174576.
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. ISSN2397-334X. PMID30833757.
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. PMID18952499.
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, PMID21295519
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. ISSN1055-7903. PMID30318266.
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)
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