View source for Cercozoa

{{Short description|Group of single-celled organisms}}
{{Automatic taxobox
| image = Cercomonas sp.jpg
| image_caption = ''[[Cercomonas]]''
| taxon = Cercozoa
| authority = [[Cavalier-Smith]], 1998<ref name="Cavalier6"/> emend. Adl et al., 2005 emend. Cavalier-Smith, 2018<ref name="ThomCav2018"/>
| synonyms=
*[[Rhizopoda]] {{au|[[Félix Dujardin|Dujardin]], 1835 ''stat. nov.'' [[Haeckel]], 1866 ''emend.'' [[Cavalier-Smith]], 1996}}<ref name="amoeboflagellates"/>
*[[Filosa]] {{au|Cavalier-Smith & Chao, 2003}}
| subdivision_ranks = Classes
| subdivision =
*[[Monadofilosa]]
**[[Eoglissa]]
***[[Metromonadea]]
***[[Helkesea]]
**[[Ventrifilosa]]
***[[Thecofilosea]]
***[[Imbricatea]]
***[[Sarcomonadea]]
*[[Reticulofilosa]]
**[[Chlorarachnea]]
**[[Granofilosea]]
**[[Skiomonadea]]?
}}

'''Cercozoa''' (now synonymised with '''Filosa''')<ref name="ThomCav2018"/> is a phylum of diverse single-celled [[eukaryote]]s.<ref name="pmid15148395"/><ref name="Ebriidphylogeny"/> They lack shared morphological characteristics at the microscopic level,<ref name="Comparative interstitial cercozoa"/> and are instead united by [[phylogeny|molecular phylogenies]] of [[rRNA]] and [[actin]] or [[Ubiquitin#Polyubiquitin chains|polyubiquitin]].<ref name="urlCERCOZOAE"/> They were the first major [[eukaryotic]] group to be recognized mainly through [[phylogeny|molecular phylogenies]].<ref name="HighDiversityCercozoa"/> They are the natural predators of many species of bacteria. They are closely related to the phylum [[Retaria]], comprising amoeboids that usually have complex shells, and together form a supergroup called [[Rhizaria]].<ref name="ThomCav2018"/>

==Characteristics==
The group includes most [[amoeboid]]s and [[flagellate]]s that feed by means of filose pseudopods. These may be restricted to part of the cell surface, but there is never a true [[cytostome]] or mouth as found in many other protozoa. They show a variety of forms<ref name="pmid14658494"/> and have proven difficult to define in terms of structural characteristics, although their unity is strongly supported by [[phylogeny|phylogenetic]] studies.

==Diversity==
Some cercozoans are grouped by whether they are "filose" or "reticulose" in the behavior of their [[cytoskeleton]] when moving:<ref name="pmid18952499"/>
*Filose, meaning their [[pseudopod]]s develop as [[filopodia]]. For example:
**[[Euglyphid]]s, filose amoebae with shells of [[silica|siliceous]] scales or plates, which are commonly found in soils, nutrient-rich waters, and on aquatic plants.
**''[[Gromia]]'', a shelled amoeba.
**[[Tectofilosid]]s, filose amoebae that produce organic shells.
**[[Cercomonad]]s, common soil-dwelling amoeboflagellates.
*Reticulose, meaning they form a reticulating net of [[pseudopod]]s. For example:
**[[Chlorarachniophyte]]s, set apart by the presence of [[chloroplast]]s bound by four membranes and still possess a vestigial nucleus, called a nucleomorph.  As such, they have been of great interest to researchers studying the endosymbiotic origins of organelles.
Other important ecological groups are:
*[[Granofilosea]], comprising several groups traditionally considered [[heliozoa]] such as [[Heliomonadida]], [[Desmothoracida]] and [[Gymnosphaerida]].<ref name="pmid18952499"/>
*[[Phaeodaria]], marine protozoa previously considered [[radiolaria]]ns.<ref name=nakamura2015/>

==Ecology==
As well as being highly diverse in [[Morphology (biology)|morphology]] and [[physiology]], Cercozoa also shows high [[ecological diversity]].<ref name="HeathlandCercozoa"/> The phylum Cercozoa includes many of the most abundant and ecologically significant [[protozoa]] in [[soil]], [[Marine life|marine]] and [[freshwater]] ecosystems.<ref name="HighDiversityCercozoa"/>

Soil-dwelling cercozoans are one of the dominant groups of free-living eukaryotic microorganisms found in [[temperate]] soils, accounting for around 30% of identifiable protozoan [[DNA]] in arid or semi-arid soils and 15% in more humid soils. In [[transcriptomic]] analyses they account for 40-60% of all identifiable protozoan [[RNA]] found in forest and grassland soils. They also comprise 9-24% of all [[operational taxonomic unit]]s found in the [[ocean floor]].<ref name="HeathlandCercozoa"/>

Some cercozoa are coprophilic or [[wikt:coprozoic|coprozoic]], meaning they use [[feces]] as a source of [[nutrient]]s or as transport through animal hosts. The faecal habitat is an understudied reservoir of microbial eukaryotic diversity, dominated by amoeboflagellates from the phylum Cercozoa. Strongly coprophilic examples of cercozoa are the flagellates ''[[Cercomonas]]'', ''[[Proleptomonas]]'' and ''[[Helkesimastix]]'', and the [[sorocarp]]ic amoeba ''[[Guttulinopsis]]''. Many new cercozoan lineages, especially among [[sarcomonad]]s, have been discovered through phylogenetic sampling of feces because they appear preferentially in this medium.<ref name="CoprophilicRhizaria"/>

Cercozoan [[bacterivore]]s (i.e. [[predation|predator]]s of [[bacteria]]) are highly diverse and important in the plant [[phyllosphere]], the leaf surfaces of plants. Particularly [[sarcomonad]]s, with their ability to [[microbial cyst|cyst]], feed and multiply within hours, are perfectly adapted to the fluctuating environmental factors in the phyllosphere. Their predation causes shifts in the bacterial communities: they reduce populations of [[alphaproteobacteria]] and [[betaproteobacteria]], which are less resistant to their grazing, in favour of other bacterial populations such as [[gammaproteobacteria]].<ref name="GrazingCercomonads"/>

==Evolution==
===External evolution===

{{cladogram|title=Paraphyletic Cercozoa<ref name="ThomCav2018"/>
|{{clade|style=font-size:90%;
|label1=[[Rhizaria]]
|1={{clade
   |label1=[[Filosa]]
   |sublabel1=(=Cercozoa)
   |1={{clade
      |1=[[Reticulofilosa]]
      |2=[[Monadofilosa]]
      }}
   |label2=[[Retaria]]
   |2={{clade
      |1=[[Endomyxa]]
      |label2=Ectoreta
      |2={{clade
         |1=[[Taxopodida]]
         |2={{clade
            |1=[[Foraminifera]]
            |2=[[Radiolaria]]
            }}
         }}
      }}
   }}
}}
}}

{{cladogram|title=Monophyletic Cercozoa<ref name="MonoCercozoa"/>
|{{clade|style=font-size:90%;
|label1=[[Rhizaria]]
|1={{clade
   |label2=[[Retaria]]
   |2={{clade
      |2=[[Acantharea]]
      |1={{clade
         |2=[[Polycystinea]]
         |1=[[Foraminifera]]
         }}
      }}
   |label1=Cercozoa
   |1={{clade
      |label1=[[Filosa]]
      |1={{clade
         |1=[[Reticulofilosa]]
         |2=[[Monadofilosa]]
         }}
      |2={{clade
         |2=''Lapot gusevi''
         |1=[[Endomyxa]]
         }}
      }}
   }}
}}
}}

Originally, Cercozoa contained both Filosa and [[Endomyxa]], according to [[phylogenetic]] analyses using [[ribosomal RNA]] and [[tubulin]]. These analyses also confirmed Cercozoa as the sister group of Retaria within the supergroup Rhizaria.<ref name="pmid18952499"/><ref name="NovelCulturedProtists"/>

However, the [[monophyly]] of the group was still uncertain. Posterior multigene phylogenetic analyses consistently found Cercozoa to be paraphyletic, because [[Endomyxa]] clustered next to [[Retaria]] instead of Filosa.<ref name="MitosomeRhizaria"/><ref name="EvolutionOfRhizaria"/><ref name="SingleCellRhizaria"/> Because of this, Endomyxa was excluded from Cercozoa, which became a synonym of Filosa.<ref name="ThomCav2018"/>

More recent phylogenomic analyses with better sampling recovered a sister relationship between Filosa (=Cercozoa) and [[Endomyxa]] once again,<ref name="MonoCercozoa"/> although the modern classification of eukaryotes retains [[Endomyxa]], Cercozoa and [[Retaria]] as separate phyla within [[Rhizaria]].<ref name="Adl 2019"/>

===Internal evolution===
The phylum Cercozoa previously contained both Filosa and [[Endomyxa]], but in the latest classifications Endomyxa has been excluded, and Cercozoa is now synonymous with Filosa. It is composed of two [[subphyla]]: [[Monadofilosa]] and [[Reticulofilosa]]. According to multigene [[phylogenetic]] analyses, Monadofilosa is a robust [[clade]], in which the deepest branching group is [[Metromonadea]], followed by [[Helkesea]] as the second group (together forming the paraphyletic [[Eoglissa]]) before the divergence of the clade [[Ventrifilosa]] ([[Imbricatea]], [[Sarcomonadea]] and [[Thecofilosea]]). On the other hand, Reticulofilosa is probably paraphyletic, with  [[Granofilosea]] diverging earlier than [[Chlorarachnea]], which makes Chlorarachnea the sister group of Monadofilosa.<ref name="ThomCav2018"/>

{{clade|style=font-size:90%;|label1='''Cercozoa'''|1={{clade|2=[[Granofilosea]]|1={{clade|2=[[Chlorarachnea]]|label1=[[Monadofilosa]]|1={{clade|2=[[Metromonadea]]|1={{clade|2=[[Helkesea]]|label1=[[Ventrifilosa]]|1={{clade|1=[[Imbricatea]]|2=[[Sarcomonadea]]|state2=double|3=[[Thecofilosea]]}}}}}}}}}}}}

A more recent [[phylogenomic]] analysis recovered both Monadofilosa and Reticulofilosa as monophyletic within the clade Filosa.<ref name="MonoCercozoa"/>

In addition to the known [[Granofilosea]], [[Chlorarachnea]] and [[Monadofilosa]], a variety of [[clades]] inside Cercozoa have been discovered in other analyses and have slowly been described and named, such as [[Tremulida]] (previously known as Novel Clade 11)<ref name="NovelCulturedProtists"/> and [[Aquavolonida]] (Novel Clade 10),<ref name="NovelClade10"/> although their specific positions among the two main cercozoan subphyla have yet to be refined. These two orders have been classified as the class [[Skiomonadea]], within Reticulofilosa.<ref name="ThomCav2018"/>

==Classification==
The classification of Cercozoa was revised in 2018:<ref name="ThomCav2018"/>
*Subphylum [[Reticulofilosa]] {{au|Cavalier-Smith 1997}}
**Class [[Chlorarachnea]] {{au|Hibberd & Norris 1984 (as [[Chlorarachniophyceae]])}}
**Class [[Granofilosea]] {{au|Cavalier-Smith & Bass 2009}}
**Class [[Skiomonadea]] {{au|Cavalier-Smith 2012}}
*Subphylum [[Monadofilosa]] {{au|Cavalier-Smith 1997}}
**Superclass [[Eoglissa]] {{au|Cavalier-Smith 2011 emend. 2018}}
***Class [[Metromonadea]] {{au|Cavalier-Smith 2007}}
***Class [[Helkesea]] {{au|Cavalier-Smith 2018}}
**Superclass [[Ventrifilosa]] {{au|Cavalier-Smith 2012 emend. 2018}}
***Class [[Sarcomonadea]] {{au|Cavalier-Smith 1993 emend. 2018}}
***Class [[Imbricatea]] {{au|Cavalier-Smith 2003 emend. 2018}}{{br}}
***Class [[Thecofilosea]] {{au|Cavalier-Smith 2003 emend. 2012}}{{br}}

==Gallery==
<gallery class="center">
File:Cercomonas sp.jpg|''Cercomonas'' sp. (Cercozoa: [[Cercomonadida]])
File:Ebria_tripartita.jpg|''Ebria'' sp. (Cercozoa: [[Ebridea]])
File:Rhipidodendron splendidum.jpg|''Rhipidodendron'' sp. (Cercozoa: [[Spongomonadea]])
File:Euglypha_sp.jpg|''Euglypha'' sp. (Cercozoa: [[Euglyphida]])
File:Haeckel Phaeodaria 1.jpg|Phaeodarians (Cercozoa: [[Phaeodarea]])
File:Clathrulina elegans - - Print - Iconographia Zoologica - Special Collections University of Amsterdam - UBAINV0274 113 04 0030.tif|''[[Clathrulina elegans]]'' (Cercozoa: [[Desmothoracida]])
File:Chlorarachnion_reptans.jpg|''[[Chlorarachnion]]'' sp. (Cercozoa: ([[Chlorarachniophyta]])
File:Vampyrella lateritia.jpg|''Vampyrella'' sp. (Cercozoa: [[Vampyrellidae]])
File:Orciraptor-gr1c.jpg|''Orciraptor agilis'' ([[Viridiraptoridae]]) attacking ''[[Mougeotia]]'' sp. ([[Zygnemataceae]])
File:Die_Gartenlaube_(1890)_b_083.jpg|''[[Gromia]]'' (Cercozoa: [[Gromiidea]])
File:Auranticordis_quadriverberis_1a.jpg|''[[Auranticordis]]'' (Cercozoa: [[Marimonadida]])
</gallery>

==References==<!-- CurrBiol17:1420. -->
{{Reflist|refs=
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<ref name="MonoCercozoa">{{Cite journal|date=2019-01-01|title=Phylogenomics supports the monophyly of the Cercozoa|journal=Molecular Phylogenetics and Evolution|language=en|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.|s2cid=52982396}}</ref>

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<ref name="NovelClade10">{{cite journal|title=Rhizarian 'Novel Clade 10' Revealed as Abundant and Diverse Planktonic and Terrestrial Flagellates, including ''Aquavolon'' n. gen.|doi=10.1111/jeu.12524|vauthors=Bass D, Tikhonenkov DV, Foster R, Dyal P, Janouškovec J, Keeling PJ, Gardner M, Neuhauser S, Hartikainen H, Mylnikov AP, Berney C|date=2018|journal=Journal of Eukaryotic Microbiology|volume=65 |issue=6 |pages=828–842 |pmid=29658156 |pmc=6282753 }}</ref>

<ref name="ThomCav2018">{{citation|first1=Thomas|last1=Cavalier-Smith|first2=Ema|last2=E. Chao|first3=Rhodri|last3=Lewis|doi=10.1007/s00709-018-1241-1|journal=Protoplasma|date=2018|volume=255|pages=1517–1574|title=Multigene phylogeny and cell evolution of chromist infrakingdom Rhizaria: contrasting cell organisation of sister phyla Cercozoa and Retaria|issue=5 |pmid=29666938 |pmc=6133090 }}</ref>

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<ref name="Ebriidphylogeny">{{cite journal|title = Ebriid phylogeny and the expansion of the Cercozoa|year=2006|last= Hoppenrath|first=M.|author2=Leander B.S.|journal = Protist|volume = 157|pages = 279–90|doi = 10.1016/j.protis.2006.03.002|pmid = 16730229|issue = 3}}</ref>

<ref name="Comparative interstitial cercozoa">Chantangsi, C. (2009). ''[https://open.library.ubc.ca/cIRcle/collections/ubctheses/24/items/1.0067363 Comparative morphology and molecular evolution of marine interstitial cercozoans.]'' PhD thesis. University of British Columbia.</ref>

<ref name="urlCERCOZOAE">{{cite web |url=http://comenius.susqu.edu/biol/202/chromalveolata/rhizariae/cercozoa/default.htm |title=SYSTEMATIC BIOLOGY: CERCOZOA |access-date=2009-03-28 |archive-date=2015-04-02 |archive-url=https://web.archive.org/web/20150402095427/http://comenius.susqu.edu/biol/202/chromalveolata/rhizariae/cercozoa/default.htm |url-status=dead }}</ref>

<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>

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<ref name="CoprophilicRhizaria">{{cite journal|title=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|doi=10.1111/1462-2920.13235|doi-access=|vauthors=Bass D, Silberman JD, Brown MW, Pearce RA, Tice AK, Jousset A, Geisen S, Hartikainen H|date=23 February 2016|journal=Environmental Microbiology|pages=1604–1619|volume=18|issue=5|pmid=26914587 }}</ref>

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<ref name="HeathlandCercozoa">{{cite journal|vauthors=Harder C, Rønn R, Brejnrod A, etal.|title=Local diversity of heathland Cercozoa explored by in-depth sequencing|journal=The ISME Journal|volume=10|pages=2488–2497|date=8 March 2016|issue=10 |doi=10.1038/ismej.2016.31|pmid=26953604 |pmc=5030685 |doi-access=free}}</ref>

<ref name="GrazingCercomonads">{{cite journal|doi=10.1111/1462-2920.13824|title=Grazing of leaf-associated Cercomonads (Protists: Rhizaria: Cercozoa) structures bacterial community composition and function|vauthors=Flues S, Bass D, Bonkowski M|journal=Environmental Microbiology|volume=19|issue=8|date=15 June 2017|pages=3297–3309|pmid=28618206 |doi-access=}}</ref>

<ref name="Adl 2019">{{cite journal|vauthors=Adl 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|year=2019|title=Revisions to the Classification, Nomenclature, and Diversity of Eukaryotes|journal=Journal of Eukaryotic Microbiology|volume=66|issue=1 |pages=4–119|doi=10.1111/jeu.12691|pmid=30257078 |pmc=6492006 }}</ref>

}}

==External links==
* [http://tolweb.org/Cercozoa/121187 Tree of Life] Cercozoa
* [https://web.archive.org/web/20160617080549/http://www.hucc.hokudai.ac.jp/~h21292/students/565.pdf phylogeny of Phaeodarea]

{{Eukaryota classification}}
{{Rhizaria}}
{{Life on Earth}}
{{Taxonbar|from1=Q500497|from2=Q1415459}}

[[Category:Cercozoa| ]]
[[Category:Protist phyla]]
[[Category:Taxa named by Thomas Cavalier-Smith]]