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{{short description|Phylum of Excavata}}
{{Automatic taxobox
| image = Naegleria (formes) en.png
| image_caption =
| taxon = Percolozoa
| display_parents = 4
| authority = [[Thomas Cavalier-Smith|Cavalier-Smith]] 1991
| subdivision_ranks = Classes
| subdivision =
*
* [[Lunosea]]
* [[Pharyngomonadea]]
|synonyms =
* Acrasiomycota <small>Whittaker 1969</small>
* Amoeboflagellates
* Pseudociliata <small>Cavalier-Smith 1993</small>
* Schizopyrenia
* Tetramitozoa <small>Dillon 1963</small>
* Stephanopogonophyta <small>Doweld 2001</small>
* Stephanopogonomorpha <small>Sleigh et al. 1984</small>
}}
The '''Percolozoa''' are a group of colourless, non-photosynthetic [[Excavata]], including many that can transform between [[amoeboid]], [[flagellate]], and [[microbial cyst|cyst]] stages.
==Characteristics==
{{unsourced|section|date=June 2024}}
Most Percolozoa are found as bacterivores in soil, fresh water and occasionally in the ocean. The only member of this group that is infectious to humans is ''[[Naegleria fowleri]]'', the causative agent of the often fatal disease [[amoebic meningitis]]. The group is closely related to the [[Euglenozoa]], and share with them the unusual characteristic of having [[mitochondrion|mitochondria]] with discoid [[crista]]e. The presence of a ventral feeding groove in the flagellate stage, as well as other features, suggests that they are part of the [[Excavata]] group.
The amoeboid stage is roughly cylindrical, typically around [[1 E-5 m|
Usually, the amoeboid form is taken when food is plentiful, and the flagellate form is used for rapid locomotion. However, not all members are able to assume both forms. The genera ''[[Percolomonas]]'', ''[[Lyromonas]]'', and ''[[Psalteriomonas]]'' are known only as flagellates, while ''[[Vahlkampfia]]'', ''[[Pseudovahlkampfia]]'', and most [[acrasid]]s do not have flagellate stages. As mentioned above, under unfavourable conditions, the acrasids aggregate to form sporangia. These are superficially similar to the sporangia of the [[dictyostelid]]s, but the amoebae only aggregate as individuals or in small groups and do not die to form the stalk.
[[File:2023 Heterolobosean.svg|center|thumb|upright=2|{{center|'''Representation of a heterolobosean'''}}{{ordered list|Eruptive [[Pseudopodia|pseudopod]]|
[[Ectoplasm (cell biology)|Ectoplasm]]|
Endoplasm|
[[Phagosome]] with prey|
[[Mitochondria|Mitochondrion]], creates [[Adenosine triphosphate|ATP]] (energy) for the cell (distoid cristae)|
[[Lysosome]], holds enzymes|
[[Food vacuole|Digestive vacuoles]] with prey|
Smooth [[Endoplasmic reticulum|endoplasmic reticulum]], the transport network for molecules going to specific parts of the cell|
[[Golgi apparatus]], modifies [[protein]]s and sends them out of the cell (reduced)|
[[Nucleolus]]|
[[Nucleus]]|
Rough endoplasmic reticulum|
[[Contractile vacuole]], regulates the quantity of water inside a cell|
Uroid}}]]
==Terminology and classification==
These are collectively referred to as schizopyrenids, amoeboflagellates, or vahlkampfids. They also include the acrasids, a group of social amoebae that aggregate to form [[sporangium|sporangia]]. The entire group is usually called the '''Heterolobosea''', but this may be restricted to members with amoeboid stages.
One Heterolobosea classification system is:<ref name="urlHeterolobosea">{{cite web |url=http://tolweb.org/Heterolobosea/96360 |title=Heterolobosea |
* Order [[Schizopyrenida]]
** Family [[Vahlkampfiidae]]
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* Order [[Acrasida]]
** Family [[Acrasidae]]
''Pleurostomum flabellatum'' has recently been added to Heterolobosea.<ref name="pmid17576098">{{cite journal |vauthors=Park JS, Simpson AG, Lee WJ, Cho BC |title=Ultrastructure and phylogenetic placement within Heterolobosea of the previously unclassified, extremely halophilic heterotrophic flagellate ''Pleurostomum flabellatum'' (Ruinen 1938) |journal=Protist |volume=158 |issue=3 |pages=397–413 |date=July 2007 |pmid=17576098 |doi=10.1016/j.protis.2007.03.004
===Phylogeny===
Based on the cladogram from Tolweb and updated
{{clade| style=font-size:90%;line-height:80%
|1={{Clade
|label1=
|1={{Clade
|label1=Pharyngomonadea
|1={{Clade
|label1=Pharyngomonadida
|1={{Clade
|label1=Pharyngomonadidae
|1=''[[Pharyngomonas]]''
}}
}}
}}
|label2=
|2={{Clade
|1={{Clade
|label1=Lunosea
|1={{Clade
|label1=Selenaionida
|1={{Clade
|label1=Selenaionidae
|1={{Clade
|1=''[[Selenaion]]''
|2=''[[Dactylomonas]]''
}}
}}
}}
}}
|2={{Clade
|label1=
|1=
|label1=Neovahlkampfida
|1={{Clade
|label1=Neovahlkampfiidae
|1=''[[Neovahlkampfia]]''
}}
}}
|label2=Eutetramitia
|2={{Clade
|label1=
|1={{Clade
|label1=
|1={{Clade
|1={{Clade
|
|1={{Clade
|1=''[[Euplaesiobystra]]''
|2=''[[Heteramoeba]]''
}}
|label2=[[Gruberellidae]]
|2={{Clade
|1=''[[Vrihiamoeba]]''
|2={{Clade
Line 81 ⟶ 119:
}}
}}
|2={{Clade
|label1=Paravahlkampfiidae
|1={{Clade
|1={{Clade
|1=''[[Fumarolamoeba]]''
|2=''[[Parafumarolamoeba]]''
}}
|2=''[[Paravahlkampfia]]''
}}
|label2=Psalteriomonadidae
|2={{Clade
|1=
|1=''[[Sawyeria]]''
|2=''[[Psalteriomonas]]''
}}
|2={{Clade
|1=
|2={{Clade
|1=''[[Pseudomastigamoeba]]''
Line 105 ⟶ 153:
|label1=[[Acrasida]]
|1={{Clade
|label1=[[Acrasidae]]
|1={{Clade
|1=
|
|
|
|2=''[[Acrasis]]''
}}
}}
}}
|label2=
|2={{Clade
|1={{Clade
|2={{Clade
|
|1={{Clade
|1=''[[Pleurostomum]]''
|2=''[[Tulamoeba]]''
}}
|label2=Naegleriidae
|2={{Clade
|1=''[[Naegleria]]''
Line 129 ⟶ 183:
}}
}}
}}
}}
|2={{Clade
|1={{Clade
|label1=[[Vahlkampfiidae]]
|1={{Clade
|1=''[[Tetramitus]]''
|2=''[[Vahlkampfia]]''
}}
}}
}}
|2={{Clade
|
|
|label1=Creneidae
|1=''[[Creneis (genus)|Creneis]]''
}}
|label2=[[Percolatea|Percolomonadida]]<ref name="Hohlfeld_2023"/>
|2={{Clade
|1={{Clade
|label1=[[Stephanopogonidae]]
|1=''[[Stephanopogon]]''
}}
|2={{Clade
|1={{Clade
|label1=Barbellidae
|1={{Clade
|1=''[[Barbelia]]''
|2=''[[Nonamonas]]''
}}
}}
|2={{Clade
|label1=Lulaidae
|1=''[[Lula (protist)|Lula]]''
|label2=Percolomonadidae
|2={{Clade
|1=''[[Percolomonas]]''
|2=''[[Nakurumonas]]''
}}
}}
}}
}}
}}
}}
Line 151 ⟶ 238:
===Taxonomy===
Phylum Percolozoa <small>Cavalier-Smith 1991</small><ref>{{cite journal | website=
* Subphylum
** Class
*** Order
**** Family
***** Genus ''[[Pharyngomonas]]'' <small>Cavalier-Smith 2008</small> [''
* Subphylum
** Genus ?''[[Costiopsis]]'' <small>Senn 1900</small>
** Genus ?''[[
** Genus ?''[[Planiosculum]]'' <small>Szabados 1948</small>
** Genus ?''[[Protomyxomyces]]'' <small>Cunningham 1881</small>
Line 168 ⟶ 255:
** Genus ?''[[Trimastigamoeba]]'' <small>Whitmore 1911</small>
** Genus ?''[[Wasielewskia]]'' <small>Hartmann & Schuessler 1913</small>
** Family Euhyperamoebidae <small>Goodkov & Seravin 1984</small> [Hyperamoebidae <small>Goodkov, Seravin & Railkin 1982</small>]
***
** Class Lunosea <small>Cavalier-Smith 2021</small>
*** Order
**** Family
***** Genus ''[[Selenaion
***** Genus ''[[Dactylomonas]]'' <small>Hanousková, Táborský & Čepička 2018</small>
** Class Neovahlkampfiea <small>Cavalier-Smith 2021</small>
*** Order Neovahlkampfiida <small>Cavalier-Smith 2021</small>
****
*****
** Class Lyromonadea <small>Cavalier-Smith 1993</small>
*** Order Lyromonadida <small>Cavalier-Smith 1993</small>
**** Family [[Gruberellidae]] <small>Page & Blanton 1985</small>
***** Genus ''[[Gruberella]]'' <small>Page 1984 non Gruber 1889 non Corliss 1960</small>
***** Genus ''[[Oramoeba]]'' <small>de Jonckheere et al. 2011</small>
***** Genus ''[[Stachyamoeba]]'' <small>Page 1975</small>
***** Genus ''[[Vrihiamoeba]]'' <small>Murase, Kawasak & Jonckheere 2010</small>
**** Family Paravahlkampfiidae
***** Genus ''[[Fumarolamoeba]]'' <small>De Jonckheere, Murase & Opperdoes 2011</small>
***** Genus ''[[Parafumarolamoeba]]'' <small>Geisen et al. 2015</small>
***** Genus ''[[Paravahlkampfia]]'' <small>Brown & de Jonckheere 1999</small>
**** Family Plaesiobystridae
***** Genus ''[[Euplaesiobystra]]'' <small>Park et al. 2009</small>
***** Genus ''[[Heteramoeba]]'' <small>Droop 1962</small>
*****
**** Family Psalteriomonadidae <small>Cavalier-Smith 1993</small> [Lyromonadidae <small>Cavalier-Smith 1993</small>]
***** Genus ''[[Harpagon]]'' <small>Panek et al. 2012</small>
***** Genus ''[[Monopylocystis]]'' <small>O'Kelly et al. 2003</small>
***** Genus ''[[Psalteriomonas]]'' <small>Broers et al. 1990</small> [''[[Lyromonas]]'' <small>Cavalier-Smith 1993</small>]
***** Genus ''[[Pseudoharpagon]]'' <small>Panek et al. 2012</small>
***** Genus ''[[Pseudomastigamoeba]]''
***** Genus ''[[Sawyeria]]'' <small>O'Kelly et al. 2003</small>
** Class [[Heterolobosea]] <small>Page & Blanton 1985</small>
*** Order [[Acrasida]] <small>Schröter 1886</small>
****
***** Genus ''[[Acrasis]]'' <small>van Tieghem 1880</small>
***** Genus ''[[
*****
***** Genus ''[[
****
***** Genus ''[[Aurem]]'' <small>Jhin & Park 2018</small>
***** Genus ''[[Pleurostomum]]'' <small>Namyslowski 1913</small>
***** Genus ''[[Tulamoeba]]'' <small>Park et al. 2009</small>
**** Family Naegleriidae <small>Kudo 1954</small> [Schizopyrenidae <small>Singh 1951 ex Singh 1952</small>; Bistadiidae <small>Doflein 1916</small>]
***** Genus ''[[Naegleria]]'' <small>Aléxéieff 1912</small> [''[[Adelphamoeba]]'' <small>Napolitano, Wall & Ganz 1970</small>; ''Didascalus'' <small>Singh 1952</small>; ''Schizopyrenus'' <small>Singh 1951 ex Singh 1952</small>]
***** Genus ''[[Willaertia]]'' <small>de Jonckheere et al. 1984</small>
*** Order Tetramitida <small>Doweld 2001</small>
**** Family [[Vahlkampfiidae]] <small>Jollos 1917 s.s.</small>
***** Genus ''[[Tetramitus]]'' <small>Perty 1852</small> [''
***** Genus ''[[Vahlkampfia]]'' <small>Chatton & LaLung-Bonnaire 1912</small> [''Guttulidium'' <small>Frenzel 1892</small>]
*** Order Creneida <small>Cavalier-Smith 2021</small>
**** Family Creneidae <small>Pánek et al. 2014</small>
***** Genus ''[[Creneis (genus)|Creneis]]'' <small>Pánek et al. 2014</small>
*** Order Percolomonadida <small>Cavalier-Smith 1993</small>
**** Family Barbeliidae <small>Arndt 2023</small><ref name="Hohlfeld_2023">{{cite journal |last1=Hohlfeld |first1=Manon |last2=Meyer |first2=Claudia |last3=Schoenle |first3=Alexandra |last4=Nitsche |first4=Frank |last5=Arndt |first5=Hartmut |title=Biogeography, autecology, and phylogeny of Percolomonads based on newly described species |journal=Journal of Eukaryotic Microbiology |date=January 2023 |volume=70 |issue=1 |pages=e12930 |doi=10.1111/jeu.12930 |pmid=35712988|doi-access=free }}</ref>
***** Genus ''[[Barbelia]]'' <small>Arndt 2021</small>
***** Genus ''[[Nonamonas]]'' <small>Hohlfeld, Meyer & Arndt 2023</small>
**** Family Lulidae <small>Hohlfeld & Arndt 2023</small><ref name="Hohlfeld_2023"/>
***** Genus ''[[Lula (protist)|Lula]]'' <small>Arndt, Nitsche & Carduck 2021</small>
**** Family [[Percolomonadidae]] <small>Cavalier-Smith 1993</small> [Choanogasteraceae]
***** Genus ''[[Percolomonas]]'' <small>Fenchel & Patterson 1986</small> [''
**** Family [[Stephanopogonidae]] <small>Corliss 1961</small>
***** Genus ''[[Nakurumonas]]'' <small>Carduck, Nitsche & Arndt 2021</small>
***** Genus ''[[Stephanopogon]]'' <small>Entz 1884</small>
==History==
The Heterolobosea were first defined by [[
{{cite journal
| last = Page | first = F.C.
Line 245 ⟶ 338:
| title = The Biology of Free-living Heterotrophic Flagellates
| editor = D.J. Patterson & J. Larsen
| pages = 113–131 | isbn =9780198577478
| publisher = Oxford University Press
}}
</ref>
Cavalier-Smith maintained the Heterolobosea as a class for amoeboid forms. He has defined Percolozoa as "Heterolobosea plus [[Percolatea]] classis nov."<ref name="pmid14657102">{{cite journal |author=Cavalier-Smith T |title=The excavate protozoan phyla Metamonada Grassé emend. (Anaeromonadea, Parabasalia, Carpediemonas, Eopharyngia) and Loukozoa emend. (Jakobea, Malawimonas): their evolutionary affinities and new higher taxa |journal=Int. J. Syst. Evol. Microbiol. |volume=53 |issue=Pt 6 |pages=1741–58 |date=November 2003 |pmid=14657102 |doi= 10.1099/ijs.0.02548-0|
==References==
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{{Eukaryota classification}}
{{Excavata}}
{{Taxonbar|from=Q22094454}}
[[Category:Percolozoa| ]]
[[Category:Bikont phyla]]
[[Category:Taxa named by Thomas Cavalier-Smith]]
|