The Percolozoa are a group of colourless, non-photosynthetic Excavata, including many that can transform between amoeboid, flagellate, and cyst stages.

Percolozoa
The three different stages of N. fowleri
Scientific classification Edit this classification
Domain: Eukaryota
Clade: Diphoda
Clade: Discoba
Superphylum: Discicristata
Phylum: Percolozoa
Cavalier-Smith 1991
Classes
Synonyms
  • Acrasiomycota Whittaker 1969
  • Amoeboflagellates
  • Pseudociliata Cavalier-Smith 1993
  • Schizopyrenia
  • Tetramitozoa Dillon 1963
  • Stephanopogonophyta Doweld 2001
  • Stephanopogonomorpha Sleigh et al. 1984

Characteristics

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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 mitochondria with discoid cristae. 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 20–40 μm in length. They are traditionally considered lobose amoebae, but are not related to the others, and unlike them, do not form true lobose pseudopods. Instead, they advance by eruptive waves, where hemispherical bulges appear from the front margin of the cell, which is clear. The flagellate stage is slightly smaller, with two or four anterior flagella anterior to the feeding groove.

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 acrasids 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 dictyostelids, but the amoebae only aggregate as individuals or in small groups and do not die to form the stalk.

 
Representation of a heterolobosean
  1. Eruptive pseudopod
  2. Ectoplasm
  3. Endoplasm
  4. Phagosome with prey
  5. Mitochondrion, creates ATP (energy) for the cell (distoid cristae)
  6. Lysosome, holds enzymes
  7. Digestive vacuoles with prey
  8. Smooth endoplasmic reticulum, the transport network for molecules going to specific parts of the cell
  9. Golgi apparatus, modifies proteins and sends them out of the cell (reduced)
  10. Nucleolus
  11. Nucleus
  12. Rough endoplasmic reticulum
  13. Contractile vacuole, regulates the quantity of water inside a cell
  14. Uroid

Terminology and classification

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These are collectively referred to as schizopyrenids, amoeboflagellates, or vahlkampfids. They also include the acrasids, a group of social amoebae that aggregate to form sporangia. The entire group is usually called the Heterolobosea, but this may be restricted to members with amoeboid stages.

One Heterolobosea classification system is:[1]

Pleurostomum flabellatum has recently been added to Heterolobosea.[2]

Phylogeny

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Based on the cladogram from Tolweb and updated by Pánek and Čepička 2014.[3][4]

Pharyngomonada
Pharyngomonadea
Pharyngomonadida
Pharyngomonadidae

Pharyngomonas

Tetramitia
Lunosea
Selenaionida
Selenaionidae
Neovahlkampfea
Neovahlkampfida
Neovahlkampfiidae

Neovahlkampfia

Eutetramitia

Taxonomy

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Phylum Percolozoa Cavalier-Smith 1991[6]

  • Subphylum Pharyngomonada Cavalier-Smith 1991
    • Class Pharyngomonadea Cavalier-Smith 2008 [Macropharyngomonadidea]
      • Order Pharyngomonadida Cavalier-Smith 2008 [Macropharyngomonadida]
        • Family Pharyngomonadidae Cavalier-Smith 2008 [Macropharyngomonadidae Cavalier-Smith 2008]
          • Genus Pharyngomonas Cavalier-Smith 2008 [Macropharyngomonas nomen nudum]
  • Subphylum Tetramitia Cavalier-Smith 1993 em. Cavalier-Smith 2008
    • Genus ?Costiopsis Senn 1900
    • Genus ?Hoehnemastix Skvortzov 1974
    • Genus ?Planiosculum Szabados 1948
    • Genus ?Protomyxomyces Cunningham 1881
    • Genus ?Protonaegleria Michel & Raether 1985
    • Genus ?Pseudovahlkampfia Sawyer 1980
    • Genus ?Schizamoeba Davis 1926
    • Genus ?Tetramastigamoeba Singh & Hanumaiah 1977
    • Genus ?Trimastigamoeba Whitmore 1911
    • Genus ?Wasielewskia Hartmann & Schuessler 1913
    • Family Euhyperamoebidae Goodkov & Seravin 1984 [Hyperamoebidae Goodkov, Seravin & Railkin 1982]
    • Class Lunosea Cavalier-Smith 2021
      • Order Selenaionida Hanousková, Táborský & Čepička 2018
        • Family Selenaionidae Hanousková, Táborský & Čepička 2018
          • Genus Selenaion Park, De Jonckheere & Simpson 2012
          • Genus Dactylomonas Hanousková, Táborský & Čepička 2018
    • Class Neovahlkampfiea Cavalier-Smith 2021
      • Order Neovahlkampfiida Cavalier-Smith 2021
        • Family Neovahlkampfiidae Hanousková, Táborský & Čepička 2018
    • Class Lyromonadea Cavalier-Smith 1993
    • Class Heterolobosea Page & Blanton 1985
      • Order Acrasida Schröter 1886
      • Order Naegleriida Starobogatov 1980
        • Genus Marinamoeba De Jonckheere et al. 2009
        • Family Tulamoebidae Kirby et al. 2015
        • Family Naegleriidae Kudo 1954 [Schizopyrenidae Singh 1951 ex Singh 1952; Bistadiidae Doflein 1916]
          • Genus Naegleria Aléxéieff 1912 [Adelphamoeba Napolitano, Wall & Ganz 1970; Didascalus Singh 1952; Schizopyrenus Singh 1951 ex Singh 1952]
          • Genus Willaertia de Jonckheere et al. 1984
      • Order Tetramitida Doweld 2001
      • Order Creneida Cavalier-Smith 2021
        • Family Creneidae Pánek et al. 2014
      • Order Percolomonadida Cavalier-Smith 1993

History

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The Heterolobosea were first defined by Page and Blanton in 1985[7] as a class of amoebae, and so only included those forms with amoeboid stages. Cavalier-Smith created the phylum Percolozoa for the extended group, together with the enigmatic flagellate Stephanopogon.[8]

Cavalier-Smith maintained the Heterolobosea as a class for amoeboid forms. He has defined Percolozoa as "Heterolobosea plus Percolatea classis nov."[9]

References

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  1. ^ "Heterolobosea". Retrieved 2009-03-25.
  2. ^ Park JS, Simpson AG, Lee WJ, Cho BC (July 2007). "Ultrastructure and phylogenetic placement within Heterolobosea of the previously unclassified, extremely halophilic heterotrophic flagellate Pleurostomum flabellatum (Ruinen 1938)". Protist. 158 (3): 397–413. doi:10.1016/j.protis.2007.03.004. PMID 17576098.
  3. ^ Pánek, Tomáš; Ptáčková, Eliška; Čepička, Ivan (2014). "Survey on diversity of marine/saline anaerobic Heterolobosea (Excavata: Discoba) with description of seven new species". International Journal of Systematic and Evolutionary Microbiology. 64 (Pt 7): 2280–2304. doi:10.1099/ijs.0.063487-0. PMID 24729392.
  4. ^ Cavalier-Smith T (December 2021). "Ciliary transition zone evolution and the root of the eukaryote tree: implications for opisthokont origin and classification of kingdoms Protozoa, Plantae, and Fungi". Protoplasma. 259 (3): 487–593. doi:10.1007/s00709-021-01665-7. PMC 9010356. PMID 34940909.
  5. ^ a b c Hohlfeld, Manon; Meyer, Claudia; Schoenle, Alexandra; Nitsche, Frank; Arndt, Hartmut (January 2023). "Biogeography, autecology, and phylogeny of Percolomonads based on newly described species". Journal of Eukaryotic Microbiology. 70 (1): e12930. doi:10.1111/jeu.12930. PMID 35712988.
  6. ^ "Part 1- Virae, Prokarya, Protists, Fungi". Collection of genus-group names in a systematic arrangement. Archived from the original on 14 August 2016. Retrieved 30 June 2016.
  7. ^ Page, F.C.; R.L. Blanton (1985). "The Heterolobosea (Sarcodina: Rhizopoda), a new class uniting the Schizopyrenida and Acrasidae (Acrasida)". Protistologica. 21: 121–132.
  8. ^ Cavalier-Smith, T. (1991). "Cell diversification in heterotrophic flagellates". In D.J. Patterson & J. Larsen (ed.). The Biology of Free-living Heterotrophic Flagellates. Oxford University Press. pp. 113–131. ISBN 9780198577478.
  9. ^ Cavalier-Smith T (November 2003). "The excavate protozoan phyla Metamonada Grassé emend. (Anaeromonadea, Parabasalia, Carpediemonas, Eopharyngia) and Loukozoa emend. (Jakobea, Malawimonas): their evolutionary affinities and new higher taxa". Int. J. Syst. Evol. Microbiol. 53 (Pt 6): 1741–58. doi:10.1099/ijs.0.02548-0. PMID 14657102.
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