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Protist classification

Classification of eukaryotes From Wikipedia, the free encyclopedia

Protist classification
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A protist (/ˈproʊtɪst/) is any eukaryotic organism (one with cells containing a nucleus) that is not an animal, plant, or fungus. The protists do not form a natural group, or clade, since they exclude certain eukaryotes with whom they share a common ancestor;[a] but, like algae or invertebrates, the grouping is used for convenience. In some systems of biological classification, such as the popular five-kingdom scheme proposed by Robert Whittaker in 1969, the protists make up a kingdom called Protista, composed of "organisms which are unicellular or unicellular-colonial and which form no tissues".[1][b] In the 21st century, the classification shifted toward a two-kingdom system of protists: Chromista (containing the chromalveolate, rhizarian and hacrobian groups) and Protozoa (containing excavates and all protists more closely related to animals and fungi).[2]

Thumb
A modern non-hierarchical interpretation of the phylogeny of eukaryotes, including protists
(the gray unlabeled line next to Sar represents Telonemia)

The following groups contain protists. The clade Opisthokonta also contains the animals and the fungi, and the kingdom Archaeplastida also contains algae and plants.

Legend:

* Lack of molecular data.
† Extinct, or exclusively fossil taxon.
? Uncertain position, reserved for above-genus taxa.
(P) Paraphyletic or polyphyletic taxon.
(P?) Potentially paraphyletic or polyphyletic taxon.
(=...) Taxonomic synonym.
(...) Same taxon in a different code of nomenclature.
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Amorphea

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Obazoa

Cladogram of Obazoa based on several analyses.[3][4][5]

The clade Obazoa contains two small groups of flagellates, the breviates and the apusomonads, and the large clade Opisthokonta, which contains animals, fungi, and their closest protist relatives.[6] Under the Cavalier-Smith system, breviates and apusomonads were two classes that composed the phylum Apusozoa,[7] but this taxon is paraphyletic, as apusomonads are more closely related to opisthokonts.[3] The taxonomy of apusomonads was expanded in a 2022 phylogenetic sudy that introduced many new genera.[8]

Opisthokonta

Opisthokonts are divided into two branches: Holozoa (animal-related), containing the ichthyosporeans, pluriformeans, filastereans and choanoflagellates; and Nucletmycea or Holomycota (fungus-related), containing the nucleariids and opisthosporidians. These groups, minus opisthosporidians, were classified as different classes within the paraphyletic phylum Choanozoa in the system of Cavalier-Smith, now obsolete. Instead, Choanozoa is the name used for the clade containing choanoflagellates and animals. Opisthosporidians (aphelids, rozellids and microsporidians) are often studied as protists, but are also considered fungi by protistologists[6] and mycologists alike.[9] Both Holozoa and Nucletmycea have been proposed once as superkingdoms by a group of mycologists who classified nucleariids and fungi as kingdoms, but without any mention of holozoan kingdoms.[10]

Amoebozoa

Cladogram of the major groups of Amoebozoa based on an 824-gene phylogenetic analysis published in 2022.[20]

The phylum Amoebozoa contains around 2,400 species[21] of primarily amoeboid protists.[6] It includes a large portion of the traditional Sarcodina, the taxon uniting all amoebae. In particular, it groups naked and testate lobose amoebae (the traditional Lobosa), as well as the archamoebae and eumycetozoans (slime molds), and a few flagellates.[22] After the general 2019 revisions published by the International Society of Protistologists (ISOP),[6] there have been specific revisions to the classification of eumycetozoans[23] and testate amoebae.[24]

Phylum Amoebozoa Lühe 1913 sensu Cavalier-Smith 1998. Genera incertae sedis: Belonocystis, Boveella, Biomyxa, Corallomyxa, Gibbodiscus, Hartmannia, Malamoeba, Malpighamoeba, Oscillosignum, Pseudothecamoeba, Rhabdamoeba, Schoutedamoeba, Stereomyxa, Subulamoeba, Triaenamoeba, Unda.[6]

  • Evosea Kang et al. 2017
    • Class Cutosea Cavalier-Smith 2016, order Squamocutida Cavalier-Smith 2016
    • Class Variosea Cavalier-Smith 2004. Genera not assigned to any lower clade: Angulamoeba, Arboramoeba, Darbyshirella, Dictyamoeba, Ischnamoeba, Heliamoeba, Filamoeba, Phalansterium.[h]
      • Family Flamellidae Lahr & Katz in Lahr et al. 2011, genera Flamella, Telaepolella Lahr et al. 2012.
      • Order Protosteliida (Protosteliales) L.S.Olive & Stoian. 1966 sensu Shadwick & Spiegel in Adl et al. 2012, family Protosteliidae (Protosteliaceae) L.S.Olive & Stoian. 1966, sole genus Protostelium (incl. Planoprotostelium).
      • Order Fractovitellida Lahr et al. 2011 sensu Kang et al. 2017
        • Family Acramoebidae Smirnov, Nassonova & Cavalier-Smith 2008, sole genus Acramoeba.
        • Family Schizoplasmodiidae Shadwick & Spiegel in Adl et al. 2012, genera Ceratiomyxella, Nematostelium, Schizoplasmodium.
        • Family Soliformoviidae Lahr & Katz 2011. Genera: Soliformovum Spiegel 1995; Grellamoeba Dyková, Kostka & Pecková 2008.
      • Order Cavosteliida (Cavosteliales) Shadwick & Spiegel in Adl et al. 2012, family Cavosteliidae (Cavosteliaceae) Olive & Stoianovitch 1964, genera Cavostelium, Schizoplasmodiopsis, Tychosporium.
      • Order Holomastigida Lauterborn 1895 (incl. Artodiscida Cavalier-Smith 2013), family Multiciliidae Poche 1913 (incl. Artodiscidae Cavalier-Smith 2013), genera Artodiscus?, Multicilia.
    • Class Archamoebea Cavalier-Smith 1983 sensu Pánek et al. 2016[39]
    • Eumycetozoa Zopf 1884 sensu Kang et al. 2017 (=Mycetozoa de Bary 1859 ex Rostafinski 1873, Macromycetozoa Fiore-Donno 2010)
      • Class Dictyostelea Lister 1909 sensu Sheikh et al. 2018 (Dictyosteliomycetes Doweld 2001). Genus unassigned to any order: Synstelium. Genus incertae sedis: Coenonia.[41]
        • Order Acytosteliales S.Baldauf S.Sheikh & Thulin 2018
          • Family Acytosteliaceae Raper ex Raper & Quinlan 1958, genera Acytostelium, Rostrostelium, Heterostelium.
          • Family Cavenderiaceae S.Baldauf, S.Sheikh & Thulin 2018, sole genus Cavenderia.
        • Order Dictyosteliales L.S.Olive ex P.M.Kirk et al. 2001. Genus unassigned to any family: Coremiostelium.
          • Family Dictyosteliaceae Rostaf. ex Cooke 1877, genera Dictyostelium, Polysphondylium.
          • Family Raperosteliaceae S.Baldauf S.Sheikh & Thulin 2018, genera Hagiwaraea, Raperostelium, Speleostelium, Tieghemostelium.
      • Class Ceratiomyxomycetes (Ceratiomyxea) D. Hawksw., B. Sutton & Ainsw. 2019,[23] order Protosporangiida Shadwick & Spiegel in Adl et al. 2012 (Ceratiomyxales G.W. Martin ex M.L. Farr & Alexop.)
        • Family Ceratiomyxaceae (Ceratiomyxidae) J.Schröt. 1889, sole genus Ceratiomyxa.
        • Family Protosporangiidae Spiegel in Adl et al. 2012 (Protosporangiaceae Leontyev, Stephenson, Schnittler, Shchepin & Novozhilov 2020), genera Clastostelium, Protosporangium.[9]
      • Class Myxogastrea Cavalier-Smith 1993 (Myxomycetes G.Winter 1880) (=Myxogastres Fr. 1829, Myxogasteres Schröt., 1889, Endosporeae Rostaf. 1873, Myxogastromycetidae G.W.Martin 1961, Myxogastromycetes M.Locquin 1974, Myxogastria Cavalier-Smith 2012 as "Myxogastria L.S.Olive 1970" nom. inval.)[23]
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Archaeplastida

  • Division Glaucophyta Skuja 1954 (=Glaucocystaceae West 1904, Glaucocystophyta Kies and Kremer 1986, Glaucoplantae Marin & Melkonian in Li et al. 2020). Incertae sedis genera: Archaeopsis, Chalarodora, Glaucocystopsis, Peliaina, Strobilomonas.[43]
  • Phylum Picozoa Seenivasan, Sausen, Medlin & Melkonian 2013 (=picobiliphytes Not et al. 2007) → class Picomonadea Seenivasan, Sausen, Medlin & Melkonian 2013 → order Picomonadida Seenivasan, Sausen, Medlin & Melkonian 2013 → family Picomonadidae Seenivasan, Sausen, Medlin & Melkonian 2013. Sole genus: Picomonas Seenivasan, Sausen, Medlin & Melkonian 2013.[i]
  • Phylum Rhodelphidia Tikhonenkov, Gawryluk, Mylnikov & Keeling 2019 → class Rhodelphidea Tikhonenkov, Gawryluk, Mylnikov & Keeling 2019 → order Rhodelphida Tikhonenkov, Gawryluk, Mylnikov & Keeling 2019 → family Rhodelphidae Tikhonenkov, Gawryluk, Mylnikov & Keeling 2019. Sole genus: Rhodelphis Tikhonenkov, Gawryluk, Mylnikov & Keeling 2019.[45]
  • Division Rhodophyta Wettstein 1901 (=Rhodophyceae Thuret 1855, emend. Rabenhorst 1863, Rhodoplantae Saunders and Hommersand 2004 emend. Adl et al. 2005)

Chloroplastida

  • Division Prasinodermophyta Marin & Melkonian in Li et al. 2020[46]
    • Class Prasinodermophyceae Marin & Melkonian in Li et al. 2020 → order Prasinodermatales Marin & Melkonian in Li et al. 2020 → family Prasinodermataceae Marin & Melkonian in Li et al. 2020. Sole genus: Prasinoderma Hasegawa & Chihara in Hasegawa et al. 1996.
    • Class Palmophyllophyceae Leliaert et al. 2016 emend. Marin & Melkonian in Li et al. 2020
      • Order Palmophyllales Zechman, Verbruggen, Leliaert, Ashwort, M.A.Buchheim, Fawley, H.Spalding, Pueschel, J.A.Buschheim, Verghese & Hanisak 2010 → family Palmophyllaceae Zechman, Verbruggen, Leliaert, Ashwort, M.A.Buchheim, Fawley, H.Spalding, Pueschel, J.A.Buschheim, Verghese & Hanisak 2010. Genera: Palmophyllum Kützing 1849; Verdigellas Ballantine & Norris 1994; Palmoclathrus? Womersley 1971.[47]
      • Order Prasinococcales Guillou et al. ex Leliaert et al. 2016 emend. Marin & Melkonian in Li et al. 2020 → family Prasinococcaceae Leliaert et al. 2016. Sole genus: Prasinococcus Miyashita & Chihara in Miyashita et al. 1993.
  • Division Chlorophyta Reichenbach 1828 emend. Pascher 1914 emend. Lewis & McCourt 2004
  • Division Streptophyta Bremer & Wanntorp 1981 (=Charophyta Migula 1897 emend. Karol et al. 2009)
    • Class Chlorokybophyceae Irisarri et al. 2021 → order Chlorokybales Irisarri et al. 2021 → family Chlorokybaceae Irisarri et al. 2021. Sole genus: Chlorokybus Geitler 1942.[55]
    • Class Mesostigmatophyceae Marin & Melkonian 1999 emend. Lewis & McCourt 2004 (=Mesostigmata Turmel et al. 2002) → order Mesostigmales Cavalier-Smith 1998 → family Mesostigmataceae Fott 1947. Sole genus: Mesostigma Lauterborn 1894.
    • Class Klebsormidiophyceae C. Jeffrey ex Guiry 2023[56]
      • Order Klebsormidiales Bierenbroodspot et al. 2024 → family Klebsormidiaceae Bierenbroodspot et al. 2024. Genera: Klebsormidium Silva, Mattox & Blackwell 1972; Interfilum Chodat 1992; Streptofilum Mikhailyuk & Lukešová 2018.
      • Order Entransiales Bierenbroodspot et al. 2024 → family Entransiaceae Bierenbroodspot et al. 2024. Sole genus: Entransia E.O. Hughes 1948.
      • Order Hormidiellales Bierenbroodspot et al. 2024 → family Hormidiellaceae Bierenbroodspot et al. 2024. Genera: Hormidiella Iyengar & Kanthamma 1940; Streptosarcina Mikhailyuk & Lukešová 2018.
    • Phragmoplastophyta Lecointre & Guyander 2006
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Pancryptista

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Cladogram of Pancryptista based on 2020 and 2022 analyses.[59][60]

The phylum Cryptista contains heterotrophic and photosynthetic single-celled flagellates. Its classification has undergone many changes through the years, and several conflicting systems and nomenclatures coexist. It was described by Thomas Cavalier-Smith in 1989 to unite two distinct groups of flagellates: the photosynthetic cryptomonads, and the heterotrophic goniomonads (respectively the two classes Cryptomonadea and Cyathomonadea). The name Cryptista was meant to be a synonym of Cryptophyta,[61] the algal division described by the same author in previous years.[62] In 2004, he modified the classification of Cryptista to add two subphyla: Cryptomonada, containing the aforementioned classes (renamed as Cryptophyceae and Goniomonadea respectively), and Leucocrypta, containing the heterotrophic katablepharids.[63] The next year, a separate group of authors proposed a different higher ranking for katablepharids as a phylum Kathablepharida or division Katablepharidophyta, related to but independent from phylum Cryptophyta, with no mention of Cryptista.[64] Between 2013 and 2015, Cavalier-Smith updated the classification once more by describing three new subphyla: Rollomonadia, containing the previous subphyla lowered to superclasses; Palpitia, containing the flagellate Palpitomonas bilix;[65] and Corbihelia, containing picozoans, telonemids, and some heliozoan species (Heliomorpha and Microheliella).[66] Corbihelia did not reach consensus: later analyses and revisions separated telonemids and picozoans as their own clades, and placed Heliomorpha in Cercozoa.[6] However, the addition of Palpitomonas and the monophyly of Rollomonadia have been supported in other analyses.[59]

Cryptista was redefined in the 2019 ISOP revision as the clade containing Palpitomonas, katablepharids, goniomonads and cryptomonads. However, this revision introduced yet another classification system that is contradictory with the previous ones. They proposed a single class Cryptophyceae uniting cryptomonads, goniomonads and katablepharids, and treated cryptomonads as a single order Cryptomonadales,[6] although this does not follow scientific consensus: there are more cryptomonad orders (e.g., Pyrenomonadales, Tetragonidiales) and the name Cryptophyceae was already used for taxa that excluded katablepharids[67] and often excluded goniomonads too.[66] There has not been a new revision since.

Unlike with Heliomorpha, the genus Microheliella was genetically sequenced and its affinities have been resolved. In 2022, it was proposed as the sister group of Cryptista, in a clade known as Pancryptista.[60]

Pancryptista Yazaki et al. 2022

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Haptista

  • Haptophyta Hibberd 1976 ex. Edvardsen & Eikrem 2000
  • Centroplasthelida Febvre-Chevalier & Febvre 1984 [Centrohelea Kühn 1926 sensu Cavalier-Smith in Yabuki et al. 2012; Centroheliozoa Dürrschmidt & Patterson 1987]
    • ?Parasphaerastrum.
    • ?Spiculophryidae Shɨshkin & Zlatogursky 2018. Spiculophrys.
    • Pterocystida Cavalier-Smith and von der Heyden 2007, emend. Shɨshkin and Zlatogursky 2018
      • Raphidista Shɨshkin & Zlatogursky 2018
        • Choanocystidae Cavalier-Smith & von der Heyden 2007. Choanocystis.
        • Raphidiophryidae Febvre-Chevalier & Febvre 1984, emend. Shɨshkin & Zlatogursky 2018. Raphidiophrys.
      • Pterista Shɨshkin & Zlatogursky 2018
        • Oxnerellidae Cavalier-Smith & Chao 2012. Oxnerella.
        • Pterocystidae Cavalier-Smith & von der Heyden 2007. Pterocystis, Raineriophrys, Chlamydaster, Pseudoraphidiophrys, Pseudoraphidocystis, Triangulopteris,[72] Khitsovia.[73]
        • Heterophryidae Poche 1913, emend. Cavalier-Smith & von der Heyden 2007. Heterophrys, Sphaerastrum.
    • Panacanthocystida Shɨshkin & Zlatogursky 2018
      • Yogsothothidae Shɨshkin & Zlatogursky 2018 [Chthonida Shɨshkin & Zlatogursky 2018; Yogsothothina Shɨshkin & Zlatogursky 2018]. Yogsothoth.
      • Acanthocystida Cavalier-Smith & von der Heyden 2007 emend. Shɨshkin & Zlatogursky 2018
        • Marophryidae Cavalier-Smith & von der Heyden 2007 emend. Shɨshkin & Zlatogursky 2018 [Marophryina Cavalier-Smith in Yabuki et al. 2012]. Marophrys.
        • Chalarothoracina Hertwig & Lesser 1874 sensu Cavalier-Smith in Yabuki et al. 2012 emend. Shɨshkin & Zlatogursky 2018
          • Acanthocystidae Claus 1874 emend. Shɨshkin & Zlatogursky 2018. Acanthocystis.
          • Raphidocystidae Zlatogursky 2018. Raphidocystis.
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Stramenopiles

  • Class Platysulcea Cavalier-Smith 2018 → order Platysulcida Cavalier-Smith 2018 → family Platysulcidae Shiratori, Nakayama & Ishida 2015. Sole genus: Platysulcus Shiratori, Nakayama & Ishida 2015.
  • Family Kaonashiidae Weston, Eglit & Simpson 2023. Sole genus: Kaonashia Weston, Eglit & Simpson 2023.[74]

Bigyra

  • Subphylum Opalozoa Cavalier Smith 1991 emend. 2006
    • Infraphylum Placidozoa Cavalier-Smith 2013
      • Class Placididea Moriya, Nakayama & Inouye 2002 → order Placidida Moriya, Nakayama & Inouye 2002 → family Placididae Moriya, Nakayama & Inouye 2002. Genera: Allegra Rybarski & Arndt 2021; Haloplacidia Rybarski, Nitsche & Arndt 2021; Pendulomonas S.M.Tong 1997; Placidia Moriya, Nakayama & Inouye 2002; Placilonga Rybarski & Arndt 2021; Suigetsumonas Okamura & Kondo 2015; Wobblia Moriya, Nakayama & Inouye 2000.[75]
      • Class Nanomonadea Cavalier-Smith in Cavalier-Smith & Scoble 2013 (=clade MAST-3) → order Uniciliatida Cavalier-Smith in Cavalier-Smith & Scoble 2013
        • Family Incisomonadidae Cavalier-Smith in Cavalier-Smith & Scoble 2013. Sole genus: Incisomonas Scoble & Cavalier-Smith 2013.
        • Family Solenicolidae Cavalier-Smith in Cavalier-Smith & Scoble 2013. Sole genus: Solenicola Pavillard 1916.
      • Class Opalomonadea Cavalier-Smith 2013 (=clade MAST-12). No described species.
      • Superclass Opalinata Wenyon 1926 emend. Cavalier-Smith 1997
        • Class Opalinea Wenyon 1926 (=Slopalinida Patterson 1985)[76]
          • Order Proteromonadida Grassé 1952 emend. Cavalier-Smith 1993 → family Proteromonadidae Grassé 1952. Sole genus: Proteromonas Kunstler 1883 (= Prowazekella Aléxéieff 1912).
          • Order Opalinida Poche 1913 stat. nov. Hall 1953 emend. Cavalier-Smith
            • Family Karotomorphidae Travis 1934. Sole genus: Karotomorpha Travis 1934.
            • Family Opalinidae Claus 1874. Genera: Cepedea Metcalf 1920; Opalina Purkinje & Valentin 1835; Protoopalina Metcalf 1918; Protozelleriella Delvinquier et al. 1991; Zelleriella Metcalf 1920.
        • Class Blastocystea Zierdt et al. 1967 → order Blastocystida Zierdt 1978 → family Blastocystidae Jiang & He 1988. Sole genus: Blastocystis Alexeieff 1911.
    • Infraphylum Bikosea Cavalier-Smith 2018 (=Bicosoecida Grassé 1926 emend. Karpov 1998) → class Bikosea Cavalier-Smith 1986 as "Bicosoecea" orth. em. 2013. Genera not assigned to families: Bilabrum Schoenle & Arndt in Schoenle et al. 2020;[77] Cafileria Jirsová, Füssy, Richtová, Gruber, & Oborník 2019;[78] Cantina Yubuki et al. 2015.
      • Family Pseudobodonidae? Cavalier-Smith in Cavalier-Smith & Scoble 2013. Sole genus: Pseudobodo Griessmann 1913.[79]
      • Subclass Rictidia Cavalier-Smith in Cavalier-Smith & Scoble 2013 → order Rictida Cavalier-Smith in Cavalier-Smith & Scoble 2013 → family Rictidae Cavalier-Smith in Cavalier-Smith & Scoble 2013. Sole genus: Rictus Yubuki, Leander & Silberman 2010.
      • Subclass BicosidiaICZN/BicosoecophycidaeICN Cavalier-Smith in Cavalier-Smith & Chao 2006
        • Superorder Borokiae Cavalier-Smith in Cavalier-Smith & Chao 2006 → order BorokidaICZN/BorokalesICN Cavalier-Smith in Cavalier-Smith & Chao 2006 → family BorokidaeICZN/BorokaceaeICN Cavalier-Smith in Cavalier-Smith & Chao 2006. Sole genus: Boroka Cavalier-Smith in Cavalier-Smith & Chao 2006.
        • Superorder Cyathobodoniae Cavalier-Smith 1993 emend. 2006. Incertae sedis genus: Halocafeteria Park, Cho & Simpson 2006.[77]
          • Order AnoecidaICZN/AnoecalesICN Cavalier-Smith 1997[l]
            • Family AnoecidaeICZN/AnoecaceaeICN Cavalier-Smith in Cavalier-Smith & Chao 2006. Sole genus: Anoeca Cavalier-Smith in Cavalier-Smith & Chao 2006.
            • Family CaecitellidaeICZN/CaecitellaceaeICN Cavalier-Smith in Cavalier-Smith & Chao 2006. Sole genus: Caecitellus Patterson et al. 1993.
            • Family CafeteriidaeICZN/CafeteriaceaeICN Moestrup 1995. Sole genus: Cafeteria Fenchel & Patterson 1988.
            • Family Symbiomonadaceae Cavalier-Smith in Cavalier-Smith & Chao 2006. Sole genus: Symbiomonas Guillou & Chretiennot-Dinet 1999.
          • Order BicoecidaICZN/BicoecalesICN Grassé 1926
            • Family Bicosoecidae Stein 1878. Genera: Bicosoeca James-Clark 1866; Stephanocodon Pascher 1942; Poteriodendron.
            • Family Labromonadidae Cavalier-Smith in Cavalier-Smith & Chao 2006. Sole genus: Labromonas Cavalier-Smith in Cavalier-Smith & Chao 2006.
          • Order PseudodendromonadidaICZN/PseudodendromonadalesICN Hibberd 1985 emend. Cavalier-Smith 2006.
            • Family PseudodendromonadidaeICZN/PseudodendromonadaceaeICN Karpov 2000. Genera: Cyathobodo Petersen & Hansen 1961; Pseudodendromonas Bourrelly 1953.
            • Family SiluaniidaeICZN/SiluaniaceaeICN Karpov 1998. Genera: Adriamonas Verhagen, Zölffel, Brugerolle & D.J.Patterson 1994; Siluania Karpov 1998; Otto Harder, Ekelund & Karpov 2014; Regin Harder, Ekelund & Karpov 2014.[80]
            • Family ParamonadidaeICZN/ParamonadaceaeICN Kent 1880. Sole genus: Paramonas Kent 1881.
            • Family NanidaeICZN/NanaceaeICN Cavalier-Smith in Cavalier-Smith & Scoble 2013. Sole genus: Nanum Kim et al. 2010 as "Nanos".
            • Family NeradidaeICZN/NeradaceaeICN Cavalier-Smith in Cavalier-Smith & Chao 2006. Sole genus: Nerada Cavalier-Smith in Cavalier-Smith & Chao 2006.
            • Family FilidaeICZN/FilaceaeICN Cavalier-Smith in Cavalier-Smith & Scoble 2013. Sole genus: Filos Kim et al. 2010.
  • Subphylum Sagenista Cavalier-Smith 1995
    • Class Labyrinthulomycetes Arx 1970 ex Dick 2001 (=Labyrinthulea L.S.Olive 1975 ex Cavalier-Smith 1989, 'phylum' LabyrinthomorphaICZN Page in Levine et al. 1980/LabyrinthulomycotaICN Whittaker 1969)[81]
      • Order Amphitremida (Poche 1913) Gomaa et al. 2013
        • Family Amphitremidae Poche 1913. Genera: Amphitrema; Archerella; Paramphitrema.
        • Family Diplophryidae Cavalier-Smith in Anderson & Cavalier-Smith 2012. Sole genus: Diplophrys Barker 1868.
      • Order Amphifilida Cavalier-Smith in Anderson & Cavalier-Smith 2012 as "Amphifiloidea"
        • Family Amphifilidae Cavalier-Smith in Anderson & Cavalier-Smith 2012. Sole genus: Amphifila Cavalier-Smith in Anderson & Cavalier-Smith 2012.
        • Family Sorodiplophryidae Cavalier-Smith in Anderson & Cavalier-Smith 2012. Genera: Fibrophrys Takahashi, Yoshida, Inouye & Watanabe 2016; Sorodiplophrys Cienkowski 1876.
      • Order OblongichytridaICZN/OblongichytrialesICN Bennett et al. 2017 → family OblongichytriidaeICZN/OblongichytriaceaeICN Cavalier-Smith in Anderson & Cavalier-Smith 2012. Sole genus: Oblongichytrium R.Yokoyama & D.Honda 2006.
      • Order LabyrinthulidaICZN Doffein 1901/LabyrinthulalesICN E.A.Bessey 1950
        • Family AplanochytriidaeICZN/AplanochytriaceaeICN Leander ex Cavalier-Smith in Anderson & Cavalier-Smith 2012. Sole genus: Aplanochytrium Bahnweg & Sparrow 1972 emend. C.A.Leander & D.Porter 2000 (=Labyrinthuloides F.O.Perkins 1973).
        • Family LabyrinthulidaeICZN Cienkowski 1867/LabyrinthulaceaeICN Haeckel 1868. Sole genus: Labyrinthula Cienkowski 1864 (=Chlamydomyxa Archer 1875, Pseudoplasmodium Molisch 1926, Labyrinthodictyon Valkanov 1969, Labyrinthorhiza Chadefaud 1956).
        • Family-level clade "StellarchytriidaeICZN/StellarchytriaceaeICN". Sole genus: Stellarchytrium FioRito & Leander 2016.
      • Order ThraustochytridaICZN/ThraustochytrialesICN Sparrow 1943
        • Family AlthorniidaeICZN/AlthornidiaceaeICN Jones & Alderman 1973. Sole genus: Althornia Jones & Alderman 1971.
        • Family ThraustochytriidaeICZN/ThraustochytriaceaeICN Sparrow ex Cejp 1959. Genera: Aurantiochytrium R.Yokoyama & D.Honda 2006; Botryochytrium R.Yokoyama, B.Salleh & D.Honda 2007; Japonochytrium Kobayasi & M.Ookubo 1953; Monorhizochytrium K.Doi & D.Honda 2017; Parietichytrium R.Yokoyama, B.Salleh & D.Honda 2007; Phycophthorum Hassett 2020;[82] Schizochytrium S.Goldst. & Belsky 1964 emend. R.Yokoyama & D.Honda 2006; Sicyoidochytrium R.Yokoyama, B.Salleh & D.Honda 2007; Thraustochytrium Sparrow 1936 emend. T.W.Johnson in Johnson & Sparrow 1961; Ulkenia A.Gaertn. 1977 emend. R.Yokoyama, B.Salleh & D.Honda 2007.
    • Class Eogyrea Cavalier-Smith 2013 (=clade MAST-6). Genera not assigned to families: Mastreximonas Lax, Cho & Keeling 2024; Vomastromonas Tikhonenkov, Prokina, Cho & Keeling 2024.[83]
      • Order Eogyrida Cavalier-Smith 2018[84] → family PseudophyllomitidaeICZN/PseudophyllomitaceaeICN Shiratori et al. 2017.[85] Sole genus: Pseudophyllomitus Lee 2002.

Gyrista

  • Subphylum Bigyromonada Cavalier-Smith 1998[86]
    • Class Developea Karpov & Aleoshin 2016 ex Cavalier-Smith 2017 → order DevelopayellidaICZN Cavalier-Smith 1987 ex 2017/DevelopayellalesICN Doweld 2001 → family DevelopayellidaeICZN/DevelopayellaceaeICN Cavalier-Smith 1997 ex 2017. Genera: Cubaremonas Tikhonenkov, Cho & Keeling 2022; Develocanicus Tikhonenkov, Cho, Mylnikov & Keeling 2022; Develocauda Tikhonenkov, Cho & Keeling 2022; Developayella Tong 1995; Develorapax Karpov & Aleoshin 2016; Mediocremonas del Campo & Weiler 2020.
    • Class Pirsonea Cavalier-Smith 2017 → order Pirsoniales Cavalier-Smith 1998 emend. Prokina et al. 2024 → Pirsoniaceae Cavalier-Smith 1998 emend. Prokina et al. 2024. Genera: Bordeauxia Prokina et al. 2024; Bullionia Prokina et al. 2024; Feodosia Prokina et al. 2024; Koktebelia Prokina et al. 2024; Noirmoutieria Prokina et al. 2024; Pirsonia Schnepf, Debres & Elbrachter 1990.[87]
  • Subphylum Pseudofungi Cavalier-Smith 1986 (=Heterokontimycotina M.W. Dick 1976, Peronosporomycotina M.W.Dick 2001)[88]: 288 
    • Class HyphochytreaICZN Cavalier-Smith 1986/HyphochytriomycetesICN Sparrow ex M.W.Dick 1983 → order Hyphochytriales Sparrow 1960
      • Family Anisolpidiaceae Karling 1943 emend. Dick 2001. Genera: Anisolpidium Karling 1943; Canteriomyces Sparrow 1960.
      • Family Hyphochytridiomycetaceae Fischer 1892 emend. Karling 1939. Sole genus: Hyphochytrium Karling 1939.
      • Family Rhizidiomycetaceae Karling 1943. Latrostium Zopf 1894; Rhizidiomyces Zopf 1884; Rhizidiomycopsis Sparrow 1960.
    • Class Oomycetes Winter 1897, emend. Dick 1976 (=Peronosporomycetes Dick 2001). Incertae sedis genera: Atkinsiella Vishniac 1958; Ciliatomyces Foissner & Foissner 1995 (=Ciliomyces Foissner & Foissner 1986);[89] Crypticola Humber et al. 1989; Ectrogella Zopf 1884; Eurychasma Magnus 1905; Halodaphnea M.W.Dick 1998; Haliphthoros Vishniac 1958; Haptoglossa Drechsler 1940; Lagena Vanterp. & Ledingham 1930; Lagenisma Drebes 1968; Olpidiopsis Cornu 1872; Pontisma H.E. Petersen 1905; Pythiella Couch 1935; Rozellopsis Karling 1942; Sirolpidium H.E.Petersen 1905.
  • Subphylum/Division Ochrophyta Cavalier-Smith 1986, emend. Cavalier-Smith & Chao 1996 (=Heterokontophyta Guiry, R.A.Andersen & Moestrup 2023). Incertae sedis genera: Commation Thomsen & Larsen 1993.[m]
    • Order Actinophryida? Claus 1874, emend. Hartmann 1926[m]
      • Suborder Actinosphaerina Cavalier-Smith 2013 → family Actinosphaeriidae Cavalier-Smith 2013. Sole genus: Actinosphaerium Ritter von Stein 1857 (=Echinosphaerium Hovasse 1965).
      • Suborder Actinophryina Cavalier-Smith 2013
        • Family Actinophryidae Dujardin 1841. Sole genus: Actinophrys Ehrenberg 1830 (=Trichoda Müller 1773 nomen oblitum, Peritricha Bory de St.Vincent 1824 nomen dubium).
        • Family Helioraphidae Cavalier-Smith 2013. Sole genus: Heliorapha Cavalier-Smith 2013.
    • Infraphylum Chrysista Cavalier-Smith 1986. Incertae sedis genera: Chrysoderma C.Billard 1984 nom. inval.; Chrysomeris Carter 1937; Chrysonephos W.R.Taylor 1952; "Giraudyopsis" Dangeard 1965 nom. inval.;[92] Rhamnochrysis Wilce & Markey 1974.[n]
      • Class Eustigmatophyceae? Hibberd & Leedale 1971[96][o]
        • Clade Goniochloridales K.P.Fawley, M.Eliáš & M.W.Fawley 2013. Genera: Goniochloris Geitler 1928; Pseudostaurastrum Chodat 1921; Tetraedriella Pascher 1930 nom. cons.; Trachydiscus H.Ettl 1964; Trebonskia Přibyl & Procházková 2022;[98] Vacuoliviride Nakayama, Nakamura, Yokoyama, Shiratori, Inouye & Ishida 2015.
        • Order Eustigmatales Hibberd 1981
          • Paraeustigmatos M.W.Fawley, Nĕmcová & K.P.Fawley 2019[99]
          • Family Eustigmataceae Hibberd 1981. Genera: Ellipsoidion Pascher 1938; Chlorobotrys Bohlin 1901; Eustigmatos Hibberd 1981; Pseudocharaciopsis K.W.Lee & Bold 1973; Vischeria Pascher 1938.
          • Family Monodopsidaceae Hibberd 1981 (includes Loboceae Hedgewald in Hedgewald, Padisak & Friedl 2007). Genera: Microchloropsis M.W.Fawley, I.Jameson & K.P.Fawley 2015; Monodopsis Hibberd 1981; Nannochloropsis Hibberd 1981; Pseudotetraedriella Hedgewald in Hedgewald, Padisak & Friedl 2007.
          • Family Neomonodaceae R.Amaral, K.P.Fawley, Nĕmcová, T.Ševčíková, Lukešová, M.W.Fawley, L.M.A.Santos & M.Eliáš 2020. Genera: Botryochloropsis? Preisig & Wilhelm 1989; Characiopsella R.Amaral, K.P.Fawley, Nĕmcová, T.Ševčíková, Lukešová, M.W.Fawley, L.M.A.Santos & M.Eliáš 2020; Munda R.Amaral, K.P.Fawley, Nĕmcová, T.Ševčíková, Lukešová, M.W.Fawley, L.M.A.Santos & M.Eliáš 2020; Neomonodus R.Amaral, K.P.Fawley, Nĕmcová, T.Ševčíková, Lukešová, M.W.Fawley, L.M.A.Santos & M.Eliáš 2020; Pseudellipsoidion (Neustupa & Nĕmcová) Nĕmcová emend. Amaral et al. 2020.
      • SII clade
        • Class Olisthodiscophyceae Barcytė, Eikrem & M. Eliáš 2021 → order Olisthodiscales Cavalier-Smith 2013 emend. Barcytė, Eikrem & M. Eliáš 2021 → family Olisthodiscaceae Cavalier-Smith 2013 emend. Barcytė, Eikrem & M. Eliáš 2021.[100] Sole genus: Olisthodiscus Carter 1937.
        • Class Pinguiophyceae Kawachi, Inouye, Honda, O'Kelly, Bailey, Bidigare & R.A.Andersen 2002 → order Pinguiochrysidales Kawachi, Inouye, Honda, O'Kelly, Bailey, Bidigare & R.A.Andersen 2002 → family Pinguiochrysidaceae Kawachi, Inouye, Honda, O'Kelly, Bailey, Bidigare & R.A.Andersen 2002. Genera: Glossomastix O'Kelly 2002; Phaeomonas Honda & Inouye 2002; Pinguiochrysis Kawachi 2002; Pinguiococcus R.A.Andersen, D.Potter & J.C.Bailey 2002; Polypodochrysis Magne 1975.
        • SSC clade[97][91]
      • SI clade[95]/PXR clade[97][91]
    • Diatomista Derelle et al. 2016, emend. Cavalier-Smith 2017 [SIII clade]. Incertae sedis taxa: environmental clades MOCH-1, MOCH-2 and MOCH-4 Massana et al. 2014.
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Alveolata

Summarize
Perspective

Palustrimonas

Ciliophora

Acavomonas

Colponema

Loeffela

Cladogram of Alveolata, based on a SSU rDNA phylogeny of basal alveolates[106] and a 203-gene phylogeny of apicomplexans and their relatives[107] published in 2023.
  • Colponemida(P) Cavalier-Smith 1993 emend. Adl et al. 2019 (=Protalveolata Cavalier-Smith 1991 emend. 2017. Genera not assigned to families: Neocolponema Gigeroff et al. 2023; Loeffela Gigeroff et al. 2023[106]
    • Phylum[p] Colponemidia Tikhonenkov et al. 2014 → class Colponemea Cavalier-Smith 1993 → order Colponemida Cavalier-Smith 1993 → family Colponemidae Cavalier-Smith & Chao 2004. Sole genus: Colponema Stein 1878.
    • Phylum[p] Acavomonidia Tikhonenkov et al. 2014 → class Acavomonea Tikhonenkov et al. 2014 → order Acavomonida Tikhonenkov et al. 2014 → family Acavomonidae Tikhonenkov et al. 2014. Acavomonas Tikhonenkov et al. 2014.
    • Order Palustrimonadida Cavalier-Smith 2017 → family Palustrimonadidae Cavalier-Smith 2017. Sole genus: Palustrimonas Patterson & Simpson 1996.

Myzozoa

Ciliophora

  • Postciliodesmatophora Gerassimova & Seravin 1976
  • Intramacronucleata Lynn 1996. Incertae sedis taxa: Protocruziidae Jankowski 1980 [Protocruziidia de Puytorac et al. 1987] (Protocruzia Faria da Cunha & Pinto 1922).
    • SAL Gentekaki et al. 2014. Incertae sedis taxa: Cariacotrichea Orsi et al. 2011 (Cariacothrix caudata), Mesodiniidae ankowski, 1980 (Mesodinium), Phacodiniidia Small and Lynn 1985 (Phacodinium).
      • Spirotrichea Bütschli 1889
        • Euplotia Jankowski 1979
          • Euplotida Small & Lynn 1985
            • Aspidiscidae Ehrenberg 1830. Aspidisca.
            • Certesiidae Borror & Hill 1995. Certesia.
            • Euplotidae Ehrenberg 1838. Euplotes.
            • Gastrocirrhidae Fauré-Fremiet 1961. Gastrocirrhus.
            • Uronychidae Jankowski 1975. Diophrys, Uronychia.
          • Discocephalida
            • Discocephalidae Jankowski 1979. Discocephalus, Prodiscocephalus, Paradiscocephalus.
            • Pseudoamphisiellidae Song et al. 1997. Leptoamphisiella, Pseudoamphisiella.
        • Perilemmaphora Berger 2008
          • Hypotrichia Stein 1859
            • Stichotrichida Fauré-Fremiet 1961 (P)[t]
              • Amphisiellidae Jankowski 1979. Amphisiella, Bistichella.
              • Atractosidae Bourland 2015. Atractos.
              • Epiclintidae Wicklow & Borror 1990. Epiclintes.
              • Gonostomatidae Small & Lynn 1985. Cotterillia, Gonostomum.
              • Halteriidae Claparède & Lachmann 1858. Halteria, Meseres.
              • Holostichidae Fauré-Fremiet 1961. Holosticha, Uncinata.
              • Kahliellidae Tuffrau 1979. Deviata, Kahliella.
              • Keronidae Dujardin 1840. Kerona.
              • Oxytrichidae Ehrenberg 1830 (P). Cyrtohymena, Gastrostyla, Oxytricha, Stylonychia.
              • Parabirojimidae Dai & Xu 2011. Parabirojimia, Tunicothrix.
              • Plagiotomidae Bütschli 1887. Plagiotoma.
              • Psammomitridae Jankowski 1979. Psammomitra.
              • Pseudoamphisiellidae Song et al. 1996. Pseudoamphisiella.
              • Psilotrichidae Bütschli 1889. Psilotricha, Urospinula.
              • Schmidingerotrichidae Foissner 2012. Schmidingerothrix.
              • Spirofilidae von Gelei 1929. Spirofilopsis, Strongylidium.
              • Trachelostylidae Small & Lynn 1985. Trachelostyla.
              • Uroleptidae Foissner & Stoeck 2008. Paruroleptus, Uroleptus.
            • Urostylida Jankowski 1979 (P)
              • Bergeriellidae Liu et al. 2010. Bergeriella, Neourostylopsis.
              • Hemicycliostylidae Lyu et al. 2018. Australothrix, Hemicycliostyla.
              • Pseudokeronopsidae Borror & Wicklow, 1983. Apoholosticha, Pseudokeronopsis.
              • Pseudourostylidae Jankowski 1979. Pseudourostyla.
              • Urostylidae Bütschli 1889. Bakuella, Diaxonella, Urostyla.
          • Oligotrichea Bütschli 1887
            • Oligotrichida Bütschli 1887
              • Cyrtostrombidiidae Agatha 2004. Cyrtostrombidium.
              • Pelagostrombidiidae Agatha 2004. Pelagostrombidium.
              • Strombidiidae Fauré-Fremiet 1970. Strombidium (P).
              • Tontoniidae Agatha 2004. Laboea, Tontonia.
            • Choreotrichida Small & Lynn 1985. Incertae sedis taxa: Lynnella Liu et al. 2011 [Lynnellidae Liu et al. 2011; Lynnellida Liu et al. 2015]
              • Strobilidiina Jankowski 1980 (P)
                • Leegaardiellidae Lynn & Montagnes 1988. Leegaardiella.
                • Lohmanniellidae Montagnes & Lynn 1991. Lohmanniella.
                • Strobilidiidae Kahl in Doflein & Reichenow 1929. Strobilidium.
                • Strombidinopsidae Small & Lynn 1985. Strombidinopsis (P).
              • Tintinnina Kofoid & Campbell 1929. Incertae sedis taxa: Helicostomella, Tintinnopsis (P), several other genera.
                • Ascampbelliellidae Corliss 1960. Ascampbelliella
                • Cyttarocylididae Kofoid & Campbell 1929 [probably synonymous with Petalotrichidae]. Cyttarocylis.
                • Dictyocystidae Haeckel 1873. Dictyocysta.
                • Epiplocylididae Kofoid & Campbell 1939. Epiplocylis.
                • Eutintinnidae Bachy et al. 2012. Dartintinnus, Eutintinnus.
                • Favellidae Kofoid & Campbell 1929. Favella.
                • Nolaclusiliidae Sniezek et al. 1991. Nolaclusilis.
                • Petalotrichidae Kofoid & Campbell 1929 [probably synonymous with Cyttarocylididae]. Petalotricha.
                • Ptychocylididae Kofoid & Campbell 1929. Cymatocylis, Ptychocylis.
                • Rhabdonellidae Kofoid & Campbell 1929. Metacylis, Rhabdonella, Schmidingerella.
                • Stenosemellidae Campbell 1954 (P). Stenosemella.
                • Tintinnidae Claparède & Lachmann 1858. Amphorellopsis, Salpingacantha, Salpingella, Tintinnus.
                • Tintinnidiidae Kofoid & Campbell 1929. Tintinnidium.
                • Undellidae Kofoid & Campbell 1929. Undella.
                • Xystonellidae Kofoid & Campbell 1929. Dadayiella, Parafavella, Xystonella.
        • Licnophoridae Bütschli 1887. Licnophora, Prolicnophora.
        • Kiitrichidae Nozawa 1941. Caryotricha, Kiitricha.
      • Lamellicorticata Vd'ačný et al. 2010
        • Armophorea Lynn 2004. Incertae sedis taxa: Mylestomatidae Kahl in Doflein & Reichenow, 1929 (Mylestoma).
          • Metopida Jankowski 1980 (P)
            • Metopidae Kahl 1927 (P). Bothrostoma, Brachonella, Eometopus, Metopus, Parametopidium, Tesnospira.
            • Apometopidae Foissner 2016. Cirranter, Urostomides.
            • Tropidoatractidae Rotterová et al. 2018. Palmarella, Tropidoatractus.
          • Clevelandellida de Puytorac & Grain 1976
            • Clevelandellidae Kidder 1938. Clevelandella.
            • Inferostomatidae Ky 1971. Inferostoma.
            • Neonyctotheridae Affa'a 1987. Neonyctotherus.
            • Nyctotheridae Amaro 1972 (P). Nyctotherus.
            • Sicuophoridae Amaro 1972. Sicuophora.
          • Caenomorphidae Poche 1913. Caenomorpha, Ludio, Sulfonecta.
          • Odontostomatida Sawaya 1940
            • Discomorphellidae Corliss 1960. Discomorphella.
            • Epalxellidae Corliss 1960. Epalxella, Saprodinium.
        • Litostomatea Small & Lynn 1981
          • Rhynchostomatia Jankowski 1980
            • Dileptida Jankowski 1978
              • Dileptidae Jankowski 1980. Apodileptus, Dileptus, Pseudomonilicaryon.
              • Dimacrocaryonidae Jankowski 1978. Dimacrocaryon, Monomacrocaryon, Rimaleptus.
              • Tracheliidae Ehrenberg 1838. Trachelius.
          • Haptoria Corliss 1974 (P). Incertae sedis taxa: Chaenea.
            • Lacrymariidae de Fromentel 1876. Lacrymaria.
            • Haptorida Corliss 1974
              • Enchelyodonidae Foissner et al. 2002. Enchelyodon, Fuscheria.
              • Homalozoonidae Jankowski 1980. Homalozoon.
              • Pleuroplitidae Foissner 1996. Pleuroplites.
            • Didiniidae Poche 1913. Didinium, Monodinium.
            • Pleurostomatida Schewiakoff 1896
              • Amphileptidae Bütschli 1889. Amphileptus.
              • Litonotidae Kent 1882. Litonotus.
              • Kentrophyllidae Wu et al. 2015. Epiphyllum, Kentrophyllum.
            • Spathidiida Foissner & Foissner 1988
              • Acropisthiidae Foissner & Foissner 1988. Acropisthium, Chaenea.
              • Actinobolinidae Kahl 1930. Actinobolina.
              • Apertospathulidae Foissner et al. 2005. Apertospathula.
              • Enchelyidae Ehrenberg 1838. Enchelys.
              • Pseudoholophryidae Berger et al. 1984. Pseudoholophrya.
              • Spathidiidae Kahl in Doflein & Reichenow 1929. Legendrea,[132] Spathidium.
              • Trachelophyllidae Kent 1882. Trachelophyllum.
            • Helicoprorodontidae Small & Lynn 1985
          • Trichostomatia Bütschli 1889
            • Vestibuliferida de Puytorac et al. 1974
              • Balantidiidae Reichenow in Doflein and Reichenow 1929. Balantidium, Neobalantidium (P).
              • Buetschliidae Poche 1913. Buetschlia.
              • Paraisotrichidae da Cunha 1917. Paraisotrichia.
              • Protocaviellidae Grain in Corliss 1979. Protocaviella.
              • Protohalliidae da Cunha & Muniz 1927. Protohallia.
              • Pycnotrichidae Poche 1913. Pycnothrix, perhaps Buxtonella.
          • Isoendo
            • Isotrichidae Bütschli 1889. Dasytricha, Isotricha.
            • Entodiniomorphida Reichenow in Doflein & Reichenow 1929
              • Blepharocorythina Wolska 1971
                • Blepharocorythidae Hsiung 1929. Blepharocorys.
                • Parentodiniidae Ito, Miyazaki & Imai 2002. Parentodinium.
                • Pseudoentodiniidae Wolska 1986. Pseudoentodinium.
              • Entodiniomorphina Reichenow in Doflein & Reichenow 1929
                • Cycloposthiidae Poche 1913. Cycloposthium.
                • Gilchristinidae Ito, Ishihara & Imai 2014 [Gilchristidae Ito et al. 2006]. Digilchristia, Gilchristina.[133]
                • Ophryoscolecidae Stein 1859. Entodinium, Ophryoscolex, Polyplastron.
                • Polydiniellidae Corliss 1960. Polydiniella.
                • Rhinozetidae Van Hoven, Gilchrist & Hamilton-Attwell 1988. Rhinozeta.
                • Spirodiniidae Strelkow 1939. Spirodinium.
                • Telamodiniidae Latteur & Dufey 1967. Telamodinium.
                • Troglodytellidae Corliss 1979. Troglodytella,.
            • Macropodiniida Lynn 2008
              • Amylovoracidae Cameron & O'Donoghue 2002. Amylovorax.
              • Macropodiniidae Dehority 1996. Macropodinium.
              • Polycostidae Cameron & O'Donoghue 2003. Polycosta.
    • CONTHREEP Lynn in Adl et al. 2012 [Ventrata Cavalier-Smith 2004]. Incertae sedis taxa: Askenasia Blochmann 1895, Cyclotrichiidae Jankowski 1980 (Cyclotrichium), Paraspathidium Noland 1937, Pseudotrachelocercidae Song 1990 (Pseudotrachelocerca), Discotrichidae Jankowski 1967 [Discotrichida Fan et al. 2014] (Discotricha, Lopezoterenia).
      • Phyllopharyngea de Puytorac et al. 1974
        • Synhymeniida de Puytorac et al. in Deroux 1978
          • Nassulopsidae Deroux in Corliss 1979. Nassulopsis.
          • Orthodonellidae Jankowski 1968. Orthodonella, Zosterodasys.
          • Scaphidiodontidae Deroux in Corliss 1979. Chilodontopsis, Scaphidiodon.
          • Synhymeniidae Jankowski in Small & Lynn 1985. Synhymenia.
        • Subkinetalia Gong et al. 2009
          • Cyrtophoria Fauré-Fremiet in Corliss 1956
            • Chlamydodontida Deroux 1976 (P)
              • Chilodonellidae Deroux 1970. Chilodonella.
              • Chitonellidae Small & Lynn 1985. Chitonella.
              • Chlamydodontidae Stein 1859. Chlamydodon.
              • Gastronautidae Deroux 1994. Gastronauta.
              • Kryoprorodontidae Alekperov & Mamajeva 1992. Gymnozoum.
              • Lynchellidae Jankowski 1968. Chlamydonella, Lynchella.
            • Dysteriida Deroux 1976
              • Dysteriidae Claparède & Lachmann 1858. Dysteria (P), Trochilia.
              • Hartmannulidae Poche 1913 (P). Hartmannula.
              • Kyaroikeidae Sniezek & Coats 1996. Kyaroikeus.
              • Plesiotrichopidae Deroux 1976. Plesiotrichopus.
              • Chonotrichia Wallengren 1895
                • Exogemmida Jankowski 1972
                  • Chilodochonidae Wallengren 1895. Chilodochona.
                  • Filichonidae Jankowski 1973. Filichona.
                  • Helichonidae Jankowski 1972. Heliochona.
                  • Lobochonidae Jankowski 1967. Lobochona.
                  • Phyllochonidae Jankowski 1972. Phyllochona.
                  • Spirochonidae Stein 1854. Spirochona.
                • Cryptogemmida Jankowski 1975
                  • Actinichonidae Jankowski 1973. Actinichona.
                  • Echinichonidae Jankowski 1973. Echinichona.
                  • Inversochonidae Jankowski 1973. Inversochona.
                  • Isochonidae Jankowski 1973. Isochona.
                  • Isochonopsidae Batisse & Crumeyrolle 1988. Isochonopsis.
                  • Stylochonidae Mohr 1948. Stylochona.
          • Rhynchodia Chatton & Lwoff 1939
            • Hypocomidae Bütschli 1889. Hypocoma.
            • Rhynchodida Chatton & Lwoff 1939
              • Ancistrocomidae Chatton & Lwoff 1939. Ancistrocoma.
              • Sphenophryidae Chatton & Lwoff 1921. Sphenophrya.
          • Suctoria Claparède & Lachmann 1858
            • Exogenida Collin 1912
              • Allantosomatidae Jankowski 1967. Allantosoma.
              • Dentacinetidae Batisse 1992. Dentacineta.
              • Dendrosomididae Jankowski 1978. Dendrosomides.
              • Ephelotidae Kent 1882. Ephelota.
              • Lecanophryidae Jankowski 1973. Lecanophrya.
              • Manuelophryidae Dovgal 2002. Manuelophrya.
              • Metacinetidae Bütschli 1889. Metacineta.
              • Ophryodendridae Stein 1867. Ophryodendron.
              • Paracinetidae Jankowski 1978. Paracineta.
              • Phalacrocleptidae Kozloff 1966. Phalacrocleptes.
              • Podophryidae Haeckel 1866. Podophrya.
              • Praethecacinetidae Dovgal 1996. Praethecacineta.
              • Rhabdophryidae Jankowski 1970. Rhabdophrya.
              • Severonidae Jankowski 1981. Severonis.
              • Spelaeophryidae Jankowski in Batisse 1975. Spelaeophrya.
              • Tachyblastonidae Grell 1950. Tachyblaston.
              • Thecacinetidae Matthes 1956. Thecacineta.
            • Endogenida Collin 1912
              • Acinetidae Stein 1859. Acineta.
              • Acinetopsidae Jankowski 1978. Acinetopsis.
              • Choanophryidae Dovgal 2002. Choanophrya.
              • Corynophryidae Jankowski 1981. Corynophrya.
              • Dactylostomatidae Jankowski 1978. Dactylostoma.
              • Dendrosomatidae Fraipont 1878. Dendrosoma.
              • Endosphaeridae Jankowski in Corliss 1979. Endosphaera.
              • Erastophryidae Jankowski 1978. Erastophrya.
              • Pseudogemmidae Jankowski 1978. Pseudogemma.
              • Rhynchetidae Jankowski 1978. Rhyncheta.
              • Solenophryidae Jankowski 1981. Solenophrya.
              • Tokophryidae Jankowski in Small & Lynn 1985. Tokophrya.
              • Trichophryidae Fraipont 1878. Trichophrya.
            • Evaginogenida Jankowski in Corliss 1979
              • Cometodendridae Jankowski 1978. Cometodendron.
              • Cyathodiniidae da Cuhna 1914. Cyathodinium.
              • Dendrocometidae Haeckel 1866. Dendrocometes.
              • Discophryidae Collin 1912. Discophrya.
              • Enchelyomorphidae Augustin & Foissner 1992. Enchelyomorpha.
              • Heliophryidae Corliss 1979. Heliophrya.
              • Periacinetidae Jankowski 1978. Periacineta.
              • Prodiscophryidae Jankowski 1978. Prodiscophrya.
              • Rhynchophryidae Jankowski 1978. Rhynchophrya.
              • Stylocometidae Jankowski 1978. Stylocometes.
              • Trypanococcidae Dovgal 2002. Trypanococcus.
      • Colpodea Small & Lynn 1981
        • Bursariomorphida Fernández-Galiano 1978
          • Bryometopidae Jankowski 1980. Bryometopus (P), Thylakidium.
          • Bursaridiidae Foissner 1993. Bursaridium, Paracondylostoma.
          • Bursariidae Bory de St. Vincent 1826. Bursaria.
        • Colpodida de Puytorac et al. 1974. Incertae sedis taxa: Bardeliellidae Foissner 1984 (Bardeliella), Hausmanniellidae Foissner 1987 (Avestina, Hausmanniella), Ilsiellidae Bourland et al. 2011 (Ilsiella), Pseudochlamydonellidae Buitkamp et al. 1989 (Hackenbergia, Pseudochlamydonella) and Marynidae Poche 1913 (Maryna).
          • Bryophryina de Puytorac et al. 1979
            • Bryophryidae de Puytorac et al. 1979. Bryophrya.
            • Sandmanniellidae Foissner & Stoeck 2009. Sandmanniella.
          • Colpodina Foissner et al. 2011
            • Colpodidae Bory de St. Vincent 1826 (P). Colpoda.
            • Grandoriidae Corliss 1960. Grandoria.
            • Tillinidae Foissner et al. 2011. Tillina.
          • Grossglockneriidae Foissner 1980 [Grossglockneriina Foissner 1980]. Grossglockneria, Pseudoplatyophrya.
        • Cyrtolophosidida Foissner 1978
          • Cyrtolophosididae Stokes 1888. Cyrtolophosis.
          • Kreyellidae Foissner 1979. Kreyella.
        • Platyophryida de Puytorac et al. 1979
          • Ottowphryidae Foissner et al. 2011. Ottowphrya, Platyophryides.
          • Platyophryidae de Puytorac et al. 1979 (P). Platyophrya.
          • Sagittariidae Grandori & Grandori 1935. Sagittaria.
          • Sorogenidae Bradbury & Olive 1980. Sorogena.
          • Woodruffiidae Gelei 1954. Etoschophrya, Rostrophrya, Woodruffia.
      • Nassophorea (P)
        • Colpodidiidae Foissner 1995. Colpodidium.
        • Nassulida Jankowski 1967
          • Furgasoniidae Corliss 1979. Furgasonia, Wolfkosia.
          • Nassulidae de Fromentel 1874. Nassula, Obertrumia.
        • Microthoracida Jankowski 1967
          • Microthoracidae Wrzesniowski 1870. Drepanomonas, Microthorax.
          • Leptopharyngidae Kahl 1926. Leptopharynx, Pseudomicrothorax.
      • Prostomatea Schewiakoff 1896 (P)
        • Apsiktratidae Foissner et al. 1994 [Prostomatida Schewiakoff 1896]
          • Balanionidae Small & Lynn 1985. Balanion.
          • Cryptocaryonidae Wright & Colorni 2002. Cryptocaryon.
          • Colepidae Ehrenberg 1838. Coleps, Plagiopogon.
          • Holophryidae Perty 1852. Holophrya.
          • Lagynidae Sola et al. 1990. Lagynus.
          • Metacystidae Kahl 1926. Metacystis, Vasicola.
          • Placidae Small & Lynn 1985. Placus.
          • Plagiocampidae Kahl 1926. Plagiocampa.
          • Prorodontidae Kent 1881. Prorodon.
          • Urotrichidae Small & Lynn 1985. Urotricha.
      • Plagiopylea Small & Lynn 1985
        • Plagiopylida Small & Lynn 1985
          • Epalxellidae Corliss 1960. Epalxella, Saprodinium.
          • Plagiopylidae Schewiakoff 1896. Plagiopyla.
          • Sonderiidae Small & Lynn 1985. Sonderia.
          • Trimyemidae Kahl 1933. Trimyema.
      • Oligohymenophorea de Puytorac et al. 1974
        • Apostomatia Chatton & Lwoff 1928
          • Apostomatida Chatton & Lwoff 1928
          • Astomatophorida Jankowski 1966
            • Opalinopsidae Hartog 1906. Opalinopsis.
          • Pilisuctorida Jankowski 1966
            • Conidophryidae Kirby 1941. Conidophrys.
        • Astomatia Schewiakoff 1896
          • Anoplophryidae Cépède 1910. Anoplophrya.
          • Buetschliellidae de Puytorac in Corliss 1979. Buetschliella.
          • Clausilocolidae de Puytorac in Corliss 1979. Clausilocola.
          • Contophryidae de Puytorac 1972. Contophyra.
          • Haptophryidae Cépède 1923. Haptophrya.
          • Hoplitophryidae Cheissin 1930. Hoplitophrya.
          • Intoshellinidae Cépède 1910. Intoshellina.
          • Maupasellidae Cépède 1910. Maupasella.
          • Radiophryidae de Puytorac 1972. Radiophrya.
        • Hymenostomatia Delage & Hérouard 1896
          • Tetrahymenida Fauré-Fremiet in Corliss 1956. Incertae sedis taxa: Trichospiridae Kahl 1926 (Trichospira).
            • Curimostomatidae Jankowski 1968. Curimostoma.
            • Glaucomidae Corliss 1971. Glaucoma.
            • Spirozonidae Kahl 1926. Spirozona.
            • Tetrahymenidae Corliss 1952. Tetrahymena.
            • Turaniellidae Didier 1971. Colpidium, Dexiostoma, Turaniella.
          • Ophryoglenida Canella 1964
            • Ichthyophthiriidae Kent 1881. Ichthyophthirius.
            • Ophryoglenidae Kent 1881. Ophryoglena.
        • Peniculia Fauré-Fremiet in Corliss 1956 (P)
          • Peniculida Fauré-Fremiet in Corliss 1956
            • Clathrostomatidae Kahl 1926. Clathrostoma.
            • Frontoniidae Kahl 1926. Disematostoma, Frontonia (P).
            • Lembadionidae Jankowski in Corliss 1979. Lembadion.
            • Maritujidae Jankowski in Small & Lynn 1985. Marituja.
            • Neobursaridiidae Dragesco & Tuffrau 1967. Neobursaridium.
            • Parameciidae Dujardin 1840. Paramecium.
            • Paranassulidae Fauré-Fremiet 1962. Paranassula.
            • Stokesiidae Roque 1961. Stokesia.
          • Urocentridae Claparède & Lachmann 1858 [Urocentrida de Puytorac, Grain & Mignot 1987]
        • Peritrichia Stein 1859
          • Sessilida Kahl 1933
            • Astylozoidae Kahl 1935. Astylozoon, Hastatella.
            • Ellobiophryidae Chatton and Lwoff 1929. Ellobiophrya.
            • Epistylididae Kahl 1933. Epistylis.
            • Lagenophryidae Bütschli 1889. Lagenophrys.
            • Operculariidae Fauré-Fremiet in Corliss 1979. Opercularia.
            • Rovinjellidae Matthes 1972. Rovinjella.
            • Scyphidiidae Kahl 1933 (P). Scyphidia.
            • Termitophryidae Lom in Corliss 1979. Termitophrya.
            • Usconophryidae Clamp 1991. Usconophrys.
            • Vaginicolidae de Fromentel 1874. Cothurnia, Pyxicola, Thuricola, Vaginicola.
            • Vorticellidae Ehrenberg 1838. Carchesium, Epicarchesium (P), Ophrydium, Pelagovorticella, Pseudovorticella (P), Vorticella (P).
            • Zoothamniidae Sommer 1951. Haplocaulus, Zoothamnium.
          • Mobilida Kahl 1933
            • Polycyclidae Poljansky 1951. Polycycla.
            • Trichodinidae Claus 1874. Trichodina.
            • Trichodinopsidae Kent 1881. Trichodinopsis.
            • Urceolariidae Dujardin 1840. Leiotrocha, Urceolaria.
        • Scuticociliatia Small 1967 (P)
          • Philasterida Small 1967
            • Cohnilembidae Kahl 1933. Cohnilembus.
            • Cryptochilidae Berger in Corliss 1979. Cryptochilum.
            • Entodiscidae Jankowski 1973. Entodiscus.
            • Entorhipidiidae Madsen 1931. Entorhipidium.
            • Orchitophryidae Cépède 1910 (P). Orchitophrya.
            • Paralembidae Corliss & de Puytorac in Small & Lynn 1985. Anophrys, Paralembus.
            • Parauronematidae Small & Lynn 1985 (P). Parauronema (P).
            • Philasteridae Kahl 1931. Kahlilembus, Philaster.
            • Pseudocohnilembidae Evans & Thompson 1964. Pseudocohnilembus.
            • Schizocaryidae Jankowski 1979. Schizocaryum.
            • Thigmophryidae Chatton & Lwoff 1926. Thigmophrya.
            • Thyrophylacidae Berger in Corliss 1961. Thyrophylax.
            • Uronematidae Thompson 1964. Uronema.
            • Urozonidae Grolière 1975. Urozona.
          • Pleuronematida Fauré-Fremiet in Corliss 1956
            • Calyptotrichidae Small & Lynn 1985. Calyptotricha.
            • Conchophthiridae Kahl in Doflein and Reichenow 1929. Conchophthirus.
            • Ctedoctematidae Small & Lynn 1985. Ctedoctema.
            • Cyclidiidae Ehrenberg 1838 (P). Cristigera, Cyclidium.
            • Dragescoidae Jankowski 1980. Dragescoa.
            • Eurystomatellidae Miao et al. 2010. Eurystomatella.
            • Histiobalantiidae de Puytorac & Corliss in Corliss 1979. Histiobalantium.
            • Peniculistomatidae Fenchel 1965. Peniculistoma.
            • Pleuronematidae Kent 1881 (P). Pleuronema.
            • Thigmocomidae Kazubski 1958. Thigmocoma.
          • Thigmotrichida Chatton & Lwoff 1922
            • Ancistridae Issel 1903. Ancistrum.
            • Hemispeiridae König 1894. Hemispeira.
            • Hysterocinetidae Diesing 1866. Hysterocineta.
            • Paraptychostomidae Ngassam et al. 1994. Paraptychostomum.
          • Loxocephalida Jankowski 1964 (P)
            • Cinetochilidae Perty 1852 (P). Cinetochilum, Sathrophilus.
            • Loxocephalidae Jankowski 1964. Cardiostomatella, Dexiotricha, Loxocephalus.
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Rhizaria

Summarize
Perspective
  • Incertae sedis taxa: Gymnosphaerida Poche 1913, emend. Mikrjukov 2000. Actinocoryne, Cienkowskya, Gymnosphaera, Hedraiophrys [possible synonym of Cienkowskya], Wagnerella.

Cercozoa

  • Incertae sedis taxa:
    • Aquavolonida Bass & Berney 2018. Aquavolon, Lapot.[134]
    • Tremulida Howe et al. 2011. Tremula.
    • Genera of uncertain placement: Dictiomyxa, Katabia, Myxodictyium, Pontomyxa, Protomyxa, Protogenes, Pseudospora, Rhizoplasma
    • Discocelia Cavalier-Smith 2013 [Discocelis Vørs 1988]
    • Psammonobiotidae* Golemansky 1974, emend. Meisterfeld 2002. Alepiella, Chardezia, Edaphonobiotus, Feuerbornia, Frenzelina, Lesquerella, Micramphora, Micropsammella, Nadinella, Ogdeniella, Psammonobiotus, Propsammonobiotus, Rhumbleriella.
  • Sarcomonadea Cavalier-Smith 1993 emend. Cavalier-Smith 2018 (P)
  • Sainouroidea Schuler et al. 2018 [Helkesida Cavalier-Smith 2018]
  • Thecofilosea Cavalier-Smith 2003, emend. Cavalier-Smith 2011. Incertae sedis taxa: Mataza.
    • Phaeodarea Haeckel 1879 [Tripylea Hertwig 1879]
      • Phaeoconchia Haeckel 1879. Coelodendrum, Coelographis, Conchellium, Conchopsis.
      • Phaeocystina Haeckel 1879. Aulacantha, Aulographis, Cannoraphis.
      • Phaeogromia Haeckel 1879. Castanella, Challengeron, Haeckeliana, Medusetta, Tuscarora.
      • Phaeosphaeria Haeckel 1879. Aulosphaera, Cannosphaera, Sagosphaera.
    • Cryomonadida Cavalier-Smith 1993
      • Rhogostomidae Dumack et al. 2017. Rhogostoma, Sacciforma, Capsellina.
      • Protaspidae Cavalier-Smith 1993 [Cryomonadidae Cavalier-Smith 1993]. Cryothecomonas, Protaspa.
    • Ventricleftida Cavalier-Smith 2011
      • Ventrifissuridae Cavalier-Smith 2018. Ventrifissura.
      • Verrucomonadidae Cavalier-Smith 2018. Verrucomonas.
    • Tectofilosida Cavalier-Smith 2003
      • Chlamydophryidae De Saedeleer, 1934. Chlamydophrys Cienkowski 1876, Diaphoropodon Archer 1869, Lecythium Hertwig & Lesser 1874, Katarium Solbach 2024,[136] Trachyrhizium Shiratori & Ishida 2016.
      • Psammonobiotidae Golemansky, 1974. Micropsamella.
      • ?Volutellidae Sudzuki, 1979. Pseudovolutella, Volutella.
      • Fiscullina Cavalier-Smith 2018
    • Ebriacea Lemmermann 1901 [Ebriidae Poche 1913; Ebriida Deflandre 1936]. Ebria, Hermesinum, Botuliforma.
  • Imbricatea Cavalier-Smith 2011
    • Spongomonadida Hibberd 1983 [Spongomonadidae Karpov 1990]. Rhipidodendron, Spongomonas.
    • Marimonadida Cavalier-Smith & Bass 2011
      • Abolliferidae Cavalier-Smith 2018. Abollifer, Cowlomonas, ?Heterochromonas.
      • Auranticordidae Cavalier-Smith 2018. Auranticordis, Rhabdamoeba.
      • Cyranomonadidae Cavalier-Smith 2018. Cyranomonas.
      • Pseudopirsoniidae Cavalier-Smith 2018. Pseudopirsonia.
    • Variglissida Cavalier-Smith 2014
      • Clautriaviidae Cavalier-Smith in Cavalier-Smith and Scoble, 2013. Clautriavia.
      • Nudifilidae Cavalier-Smith in Howe et al., 2011. Nudifila.
      • Quadriciliidae Cavalier-Smith 2018. Quadricilia.
    • Silicofilosea Adl et al. 2005, emend. Adl et al. 2012
      • Thaumatomonadida Shirkina 1987 [Thaumatomastigidae Patterson and Zoelfell 1991]
        • Thaumatomonadidae Hollande 1952. Allas, Hyaloselene, Reckertia, Thaumatomonas, Thaumatomastix, Ovaloplaca, Scutellomonas, Thaumatospina, Penardeugenia.
        • Peregriniidae Cavalier-Smith 2011. Gyromitus, Peregrinia.
        • Esquamulidae Shiratori, Yabuki & Ishida 2012. Esquamula.
      • Euglyphida Copeland 1956, emend. Cavalier-Smith 1997. Incertae sedis taxa: Phaeobola.[138]
        • Cyphoderiidae de Saedeleer 1934. Campascus*, Corythionella, Cyphoderia [Schaudinnula], Messemvriella*, Pseudocorythion, Ichthyosquama, Knarr, Oleiformis, Psammoderia.[139]
        • Paulinellidae de Saedeleer 1934, emend. Adl et al. 2012. Micropyxidiella, Ovulinata, Paulinella.
        • Euglyphina Kosakyan et al. 2016
          • Assulinidae Lara et al. 2007. Assulina, Placocista, Valkanovia*.
          • Sphenoderiidae Chatelain et al. 2013. Sphenoderia, Trachelocorythion, Deharvengia*.
          • Trinematidae Hoogenraad & De Groot 1940, emend Adl et al. 2012. Corythion, Playfairina*, Puytoracia*, Trinema.
          • Euglyphidae Wallich 1864, emend Lara et al. 2007. Euglypha, Scutiglypha.
          • Tracheleuglypha[u] Deflandre 1928
      • Trivalvulariida Siemensma & Dumack 2020.[140] Leptogromia, Trivalvularis.
  • Metromonadea Cavalier-Smith 2007, emend. Cavalier-Smith 2011. Metromonas, Metopion, Micrometopion, Kiitoksia.
  • Granofilosea Cavalier-Smith & Bass 2009.[141] Incertae sedis taxa*: ?Apogromia, ?Kibisidytes, ?Leucodictyon, ?Limnofila, ?Mesofila, ?Microcometes, ?Microgromia, ?Nanofila, ?Reticulamoeba and probably ?Belaria, ?Ditrema, ?Heliomorpha [=Dimorpha] and ?Paralieberkuehnia.
  • Chlorarachnea Hibberd & Norris 1984. Amorphochlora, Bigelowiella, Chlorarachnion, Cryptochlora, Gymnochlora, Lotharella, Minorisa, Partenskyella, Viridiuvalis.

Endomyxa

Retaria

Foraminifera

Subphylum Foraminifera d'Orbigny 1826

  • ?Lagenida
  • ?Heterogromia
  • ?Komokiacea*
  • ‘Monothalamids’(P) sensu Pawlowski et al. 2013 (Monothalamea Haeckel 1862). Genera not assigned to any lower taxon: Astrammina, Crithionina, Notodendrodes, Psammophaga, Bathysiphon, Nujappikia Gooday & Holzmann 2022.[151]
    • Family Lieberkuehniidae Siemensma et al. 2021, genera Lieberkuehnia, Claparedellus.[152]
    • Family Edaphoallogromiidae Siemensma et al. 2021, sole genus Edaphoallogromia.[152]
    • Family Lacogromiidae Siemensma & Holzmann 2021, sole genus Lacogromia.[152]
    • Family Limnogromiidae Holzmann & Siemensma 2021, sole genus Limnogromia.[152]
    • Family Reticulomyxidae Page 1987 emend. Hülsmann 2014. Reticulomyxa, Dracomyxa, Haplomyxa,[153] Wobo.
    • Family Schizamminidae Nørvang, 1961, genera Jullienella, Schizammina, Spiculosiphon.[154]
    • Family Velamentofexidae Völcker & Clauß 2021, sole genus Velamentofex.[152]
    • Order Allogromiida Loeblich & Tappan 1961
    • Order Astrorhizida Lankester 1885
  • Class Tubothalamea Pawlowski et al. 2012. †Fusulinida Delage & Hérouard 1896 probably belongs to this group.[6]
  • Class Globothalamea Pawlowski et al. 2012
    • Order Rotaliida Delage & Hérouard 1896
      • Family Planorbulinidae Schwager 1877. Planorbulinella, Hyalinea.
      • Superfamily Discorboidea Ehrenberg 1838
        • Family Discorbidae Ehrenberg 1838. Discorbis.
        • Family Rosalinidae Reiss 1963. Rupertina, Discanomalina, "Rosalina", Gavelinopsis, Planorbulina.
      • Superfamily Rotalioidea Ehrenberg 1839, emend. Pawlowski 2013. Incertae sedis taxa: ?Criboelphidium, ?"Elphidium", ?Protelphidium.
        • Family Elphidiidae Galloway 1933. Elphidium.
        • Family Ammoniidae Saidova 1981. Ammonia.
        • Family Elphidiellidae Holzmann & Pawlowski 2017. Elphidiella.
        • Family Haynesinidae Mikhalevich 2013. Haynesina, Aubignyna.
      • Superfamily Glabratelloidea Loeblich & Tappan 1964
        • Family Rotaliellidae Loeblich & Tappan 1964. Rotaliella, Rossyatella.
        • Family Buliminoididae Seiglie 1970. Buliminoides.
        • Family Glabratellidae Loeblich & Tappan 1964. Glabratella, Glabratellina, Angulodiscorbis, Planoglabratella.
      • Superfamily Calcarinoidea Schwager 1876
        • Family Calcarinidae Schwager 1876. Neorotalia, Baculogypsina, Baculogypsinoides, Schlumbergerella, Pararotalia.
      • Superfamily Nummulitoidea de Blainville 1827, family Nummulitidae de Blainville 1827. Nummulites, Operculinella, Cycloclypeus, Heterostegina, Operculina, Planoperculina, Planostegina.
      • Superfamily Serioidea Holzmann & Pawlowski 2017
        • Family Uvigerinidae Haeckel 1894. Uvigerina, Rectuvigerina, Trifarina.
        • Family Bolivinitidae Glaessner 1937. Bolivina, Brizalina, Saidovina.
        • Family Cassidulinidae d'Orbigny 1839. Globocassidulina, Cassidulinoides, Evolvocassidulina, Islandiella, Ehrenbergina
        • Family Sphaeroidinidae Cushman 1927. Sphaeroidina.
        • Family Globobuliminidae Cushman 1927. Globobulimina.
      • Incertae sedis families
        • ?Family Nonionidae Schultze 1854. Nonion, Nonionella, Nonionellina, Nonionoides.
        • ?Family Virgulinellidae Loeblich & Tappan 1984. Virgulinella.
        • ?Family Buliminidae Jones 1875. Bulimina.
        • ?Family Epistominellidae Holzmann & Pawlowski 2017. Epistominella.
        • ?Family Stainforthiidae Reiss 1963. Stainforthia, Gallitellia.
        • ?Family Cibicididae Cushman 1927. Cibicides, Cibicidoides, Heterolepa.
        • ?Family Chilostomellidae Brady 1881. Chilostomella.
        • ?Family Pullenidae Schwager 1877. Pullenia.
        • ?Family Nuttalidae Saidova 1981. Nuttalides.
        • ?Family Discorbinellidae Sigal 1952. Discorbinella, Hanzawaia.
        • ?Family Astrononionidae Cushman & Edwards 1937. Astrononion.
        • ?Family Oridorsalidae Loeblich & Tappan 1984. Oridorsalis.
        • ?Family Melonidae Holzmann & Pawlowski 2017. Melonis.
        • ?Family Cymbaloporidae Cushman 1927. Cymbaloporella.
        • ?Family Rubratelliidae Holzmann & Pawlowski 2017. Rubratella.
        • ?Family Murrayinelliidae Holzmann & Pawlowski 2017. Murrayinella.
    • Order Globigerinida Lankester 1885 emend. Arz, Arenillas & Gilabert 2022[157] (P)
      • †Superfamily Parvularugoglobigerinoidea Arenillas, Arz & Gilabert 2022
        • †Family Parvularuglobigerinidae Arenillas, Arz & Gilabert 2022. Pseudocaucasina, Palaeoglobigerina, Parvularugoglobigerina.
      • Superfamily Eoglobigerinoidea BouDagher-Fadel 2012
        • †Family Eoglobigerinidae Blow 1979. Alicantina, Eoclavatorella, Eoglobigerina, Parasubbotina, Pseudoglobigerinella, Subbotina, Turbeogloborotalia.
        • Family Globoquadrinidae Blow 1979. Dentoglobigerina, Globoquadrina.
        • †Family Neoacarininidae BouDagher-Fadel 2012. Neoacarinina.
        • †Family Porticulasphaeridae Banner 1982. Globigerinatheka, Guembelitrioides, Inordinatosphaera, Orbulinoides, Porticulasphaera.
      • Superfamily Globigerinoidea Carpenter, Parker & Jones 1862
        • Family Globigerinidae Carpenter, Parker & Jones 1862. Alloglobigerinoides, Ciperoella, Globigerina, Globicuniculus, Globigerinoides, Globigerinoidesella, Globoturborotalita, Trilobigerina [Trilobatus], Zeaglobigerina.
        • Family Globigerinellidae BouDagher-Fadel 2012. Beella, Bolliella, Globigerinella, Orcadia, Protentella, Quiltyella.
        • Family Hastigerinidae Bolli, Loeblich & Tappan 1957. Hastigerina, Hastigerinella, Hastigerinopsis.
        • Family Orbulinidae Schultze 1854. Biorbulina, Candorbulina, Orbulina, Praeorbulina.
        • Family Sphaeroidinellidae Banner & Blow 1959. Prosphaeroidinella, Sphaeroidinella, Sphaeroidinellopsis.
        • Family Turborotalitidae Hofker 1976. Berggrenia, Turborotalita.
      • Superfamily Globorotalioidea Cushman 1927
        • †Family Catapsydracidae Bolli, Loeblich & Tappan 1957. Catapsydrax, Globigerinopsoides, Globigerinoita, Velapertina.
        • Family Globorotaloididae Banner & Blow 1959. Clavatorella, Globorotaloides, Protentelloides.
        • Family Globorotaliidae Cushman 1927. Dentigloborotalia, Globoconella, Globorotalia, Hirsutella, Jenkinsella, Menardella, Neogloboquadrina, Paragloborotalia, Truncorotalia.
        • Family Pulleniatinidae Cushman 1927. Pulleniatina.
      • †Superfamily Globanomalinidoidea Loeblich & Tappan 1984, family Globanomalinidae Loeblich & Tappan 1984. Carinoturborotalia, Globanomalina, Luterbacheria, Planoglobanomalina, Turborotalia.
      • †Superfamily Hantkeninoidea Cushman 1927
        • †Family Pseudohastigerinidae Canudo 1990. Pseudohastigerina.
        • †Family Hantkeninidae Cushman 1927. Applinella, Aragonella, Clavigerinella, Cribrohantkenina, Hantkenina.
      • †Superfamily Truncorotaloidinoidea BouDagher-Fadel 2012
        • †Family Globigerapsidae Blow 1979. Globigerapsis, Muricoglobigerina.
        • †Family Planorotalitidae BouDagher-Fadel 2012. Astrorotalia, Planorotalites.
        • †Family Truncorotaloididae Loeblich & Tappan 1961. Acarinina, Igorina, Morozovella, Morozovelloides, Pearsonites, Praemurica, Pseudogloboquadrina, Testacarinata, Truncorotaloides.
    • †Order Globotruncanida Arz, Arenillas & Gilabert 2022[157]
      • †Superfamily Abathomphaloidea Arz, Arenillas & Gilabert 2022
        • †Family Abathomphalidae Pessagno 1967. Abathomphalus.
        • †Family Globotruncanellidae Maslakova 1964. Globotruncanella, Spinoglobotruncanella.
      • †Superfamily Favuselloidea Longoria 1974
        • †Family Conoglobigerinidae BouDagher-Fadel, Banner & Whittaker 1997. Conoglobigerina, Tenuigerina, ?Sphaerogerina.
        • †Family Favusellidae Longoria 1974. Ascoliella, Favusella, Koutsoukosia.
        • †Family Globuligerinidae Loeblich & Tappan 1984. Compactogerina, Globuligerina, Haeuslerina, Petaloglobigerina.
      • †Superfamily Hedbergelloidea Longoria & Gamper 1975
        • †Family Ananiidae El-Nakhal 2010. Anania, Badriella, Costellagerina, Hillsella, Liuenella [Liuella], Loeblichella, Muricohedbergella, Paracostellagerina, Pessagnoina, Planohedbergella, Pseudoclavihedbergella, Vanhintella.
        • †Family Hedbergellidae Loeblich & Tappan 1961. Asterohedbergella, Clavihedbergella, Hedbergella, Microhedbergella, Paraticinella, Pseudoguembelitria.
        • †Family Helvetoglobotruncanidae Lamolda 1976. Angulocarinella, Bermudeziana, Bollitruncana, Fingeria, Hedbergellita, Brittonella, Helvetoglobotruncana, Unitruncatus, Whiteinella.
        • †Family Praehedbergellidae Banner & Desai 1988. Blefuscuiana, Gorbachikella, Praehedbergella, Lilliputianella, Lilliputianelloides, Wondersella.
      • †Superfamily Globotruncanoidea Brotzen 1942
        • †Family Globotruncanidae Brotzen 1942. Contusotruncana, Globotruncana, Marginotruncana, Obliquacarinata, Sphaerotruncana, Ventrotruncana.
        • †Family Praeglobotruncanidae Ion 1983. Concavatotruncana, Dicarinella, Falsotruncana, Rotundina, Praeglobotruncana, Verotruncana.
        • †Family Reissidae Korchagin 2001. Elevatotruncana, Globotruncanita, Kassabiana, Radotruncana, Sigalitruncana, Turbotruncana.
      • †Superfamily Planomalinoidea Bolli, Loeblich & Tappan 1957
        • †Family Eohastigerinellidae Loeblich & Tappan 1984. Eohastigerinella, Hastigerinoides.
        • †Family Globigerinelloididae Longoria 1974. Alanlordella, Allotheca, Biglobigerinella, Blowiella, Claviblowiella, Globigerinelloides, Pseudoschackoina.
        • †Family Planomalinidae Bolli, Loeblich & Tappan 1957. Bannerina, Planomalina, Pseudoplanomalina.
        • †Family Schackoinidae Pokorný 1958. Asymetria, Groshenyia, Leupoldina, Neoschackoina, Schackoina.
      • †Superfamily Rotaliporoidea Sigal 1958
        • †Family Rotaliporidae Sigal 1958. Anaticinella, Pseudothalmanninella, Rotalipora, Thalmanninella.
        • †Family Ticinellidae Longoria 1974. Biticinella, Claviticinella, Ticinella.
      • †Superfamily Rugoglobigerinoidea Subbotina 1959
        • †Family Helvetiellidae Longoria & Gamper 1984. Archaeoglobigerina, Bucherina, Dorbignya, Edgarinella, Gandolfia, Gansserina, Globocarinata, Helvetiella, Kuglerina.
        • †Family Rugoglobigerinidae Subbotina 1959. Archaeoglobitruncana, Plummerita, Rugoglobigerina, Rugosocarinata, Rugotruncana, Trinitella.
    • †Order Heterohelicida Fursenko 1958 emend. Arz, Arenillas & Gilabert 2022[157]
      • †Superfamily Heterohelicoidea Cushman 1927
        • †Family Gublerinidae Aliyulla 1977. Gublerina, Lipsonia, Praegublerina.
        • †Family Heterohelicidae Cushman 1927. Braunella, Bronnimannella, Ehrenbergites, Globoheterohelix, Fleisherites, Hartella, Heterohelix, Huberella, Laeviheterohelix, Lazarusina, Lunatriella, Magellanina, Mihaia, Nederbragtina, Planoheterohelix, Protoheterohelix, Pseudoplanoglobulina, Steineckia, Striataella, Texasina, ?Bifarina, ?Rectoguembelina, ?Zeauvigerina.
        • †Family Pseudotextulariidae Maamouri & Salaj 1978. Planoglobulina, Pseudotextularia, Racemiguembelina.
        • †Family Pseudoguembelinidae Aliyulla 1977. Eicheriella, Leptobimodalia, Pseudoguembelina.
        • †Family Spiroplectidae Cushman 1911. Hendersonites [Hendersonia], Neohendersonites, Paraspiroplecta, Spiroplecta.
        • †Family Ventilabrellidae Maamouri & Salaj 1978. Planulitella, Proliferania, Sigalia, Ventilabrella.
    • †Order Guembelitriida Arenillas, Arz & Gilabert 2022[157]
      • †Superfamiily Cassigerinelloidea Bolli, Loeblich & Tappan 1957, family Cassigerinellidae Bolli, Loeblich & Tappan 1957. Cassigerinella, Riveroinella.
      • †Superfamily Guembelitrioidea Montanaro-Gallitelli 1957, family Guembelitriidae Montanaro-Gallitelli 1957. Cassigerinelloita, Guembelitria, Chiloguembelitria, Jenkinsina, ?Archaeoguembelitria, ?Gallitellia, ?Guembelitriella.
      • †Superfamily Chiloguembelinoidea Reiss 1963, family Chiloguembelinidae Reiss 1963. Chiloguembelina, Woodringina, ?Streptochilus.
      • †Superfamily Globoconusoidea BouDagher-Fadel 2012, family Globoconusidae BouDagher-Fadel 2012. Globoconusa, Trochoguembelitria,
    • Order ? (unnamed)[157]
      • Superfamily Globigerinitoidea BouDagher-Fadel 2012, emend. Arenillas, Arz & Gilabert 2022[157]
        • Family Candeinidae Cushman 1927. Candeina.
        • †Family Globigerinatellidae BouDagher-Fadel 2012. Globigerinatella.
        • Family Globigerinitidae Bermúdez 1961. Globigerinita, Mutabella, Polyperibola, Tinophodella.
        • Family Tenuitellidae BouDagher-Fadel 2012. Praetenuitella, Tenuitella, Tenuitellinata, Tenuitellita.
        • †Family ? (unnamed). Dipsidripella, ?Antarcticella.
    • Order Robertinida Loeblich & Tappan 1984. Hoeglundina, Robertina, Robertinoides.
    • Order Textulariida(P) Delage & Hérouard 1896. Cyclammina, Eggerella, Reophax, Textularia, Trochammina.
    • Order Carterinida Loeblich & Tappan 1981. Carterina.

Radiolaria

  • Radiolaria Müller 1858, sensu Adl et al. 2005
    • Acantharea Haeckel 1881, emend. Mikrjukov 2000
      • Chaunocanthida Schewiakoff 1926. Amphiacon, Conacon, Gigartacon, Heteracon, Stauracon.
      • Holocanthida Schewiakoff 1926. Acanthochiasma, Acanthocolla, Acanthoplegma.
      • Symphyacanthida Schewiakoff 1926. Amphilithium, Astrolonche, Pseudolithium.
      • Arthracanthida Schewiakoff 1926. Acanthometra, Daurataspis, Dictyacantha, Diploconus, Phractopelta, Phyllostaurus, Pleuraspis, Stauracantha.
    • Taxopodida Fol 1883. Sticholonche.
    • Polycystinea Ehrenberg 1839 stat. nov. Levine et al. 1980[158]
      • Spumellaria Ehrenberg 1876 stat. nov. Haeckel 1884
        • Hexacromyoidea Haeckel 1882 stat. nov. Suzuki et al. 2021 [Hexalonchata Afanasieva et al. 2005]
          • Hexacaryidae Haeckel 1882 stat. nov. Suzuki et al. 2021: Clevepilegma Dumatrica 2013, Haliphormis Ehrenberg 1846 [Hexastylanthus, Hexastylettus; Hexastylissus; Hexastylurus], Hexacaryum Haeckel 1882, Hexalonchetta Haeckel 1887, Hexancistra Haeckel 1879 [Hexancora], Hexapitys Haeckel 1882.
          • Hexacromyidae Haeckel 1882 stat. nov. Suzuki et al. 2021: ?Carpocanthum Chen & Tan 1989, Hexacromyum Haeckel 1882 [Cubosphaera; Hexacontura], Hexalonchilla Haeckel 1887 [Hexalonchusa; Staurolonchantha], Nanina Kozur & Mostler 1982 [Pentactinosphaera].
          • Hollandosphaeridae Deflandre 1973: Anomalosoma Loeblich & Tappan 1961 [Heterosoma Hollande & Enjumet 1960, Hollandosphaera Deflandre 1973 [Heliaster Hollande & Enjumet 1960], Tetrapetalon Hollande & Enjumet 1960.
        • Spongosphaeroidea Haeckel 1862 → Spongosphaeridae Haeckel 1862: Diplospongus Mast 1910, Spongodendron Hollande & Enjumet 1960, Spongosphaera Ehrenberg 1847 [Hexadoridium; Spongosphaeromma].
        • Lithocyclioidea Ehrenberg 1846
          • Astracturidae Haeckel 1882: Astromma Ehrenberg 1846 [Astractinium; Astractura; Astracturium; Astrococcura; Staurococcura], Amphactura Haeckel 1882 [?Dicoccura; ?Diplactinium], Hymeniastrum Ehrenberg 1846 [Hymenastrella; Hymenactura; Hymenacturium; Trigonactinium; Hymenactinium; Pentactura; Trigonacturium].
          • Lithocycliidae Ehrenberg 1846: Cromyatractus Haeckel 1887 [Cromyatractium; Caryatractus], Heliosestrum Haeckel 1882 [Heliosestantha; Astrosestrum; Astrosestantha; Astrophacura; Staurodiscus Krasheninnikov 1960]; Heliostylus Haeckel 1882 [Astrostylus; Stylodiscus; Stylentodiscus], Lithocyclia Ehrenberg 1846 [Astrocyclia; Coccodiscus], Phacostaurus Haeckel 1882 [Phacostaurium, Astrostaurus, Crucidiscus, Heliostaurus, Sethostaurium, Sethostaurus, Staurentodiscus], Phacotriactis Sutton 1896, Sethostylus Haeckel 1882 [Sethostylium; Amphicyclia; Phacostylus; Phacostylium], Staurocyclia Haeckel 1882 [Coccostaurus], Triactiscus Haeckel 1882 [Trigonocyclia].
          • Panartidae Haeckel 1887: Cannartus Haeckel 1882 [Cannartidissa; Cannartiscus; Pipetta; Pipettaria; Pipettella; Druppula; Druppuletta], Diartus Sanfilippo & Riedel 1980, Didymocyrtis Haeckel 1862 [Artidium; Cyphinura; Cyphocolpus; Desmartus; Ommatocampula; Panaromium; Panartus; Panartella; Panartidium; Panartissa; Panartura; Peripanartium; Peripanartula; Peripanartus; Peripanicula], Spongolivella Dumitrica 2021.
          • Phacodiscidae Haeckel 1882: Periphaena Ehrenberg 1874 [Astrophacomma; Heliodiscomma; Perizona], Phacodiscus Haeckel 1882 [Phacodiscinus; Astrophacilla; Paracenodiscus; Phacodisculus; Prunulum; Prunuletta; Sethodiscinus].
        • Spongodiscoidea Haeckel 1862 emend. Suzuki 2021
          • Spongodiscidae Haeckel 1862: Spongaster Ehrenberg 1861 [Spongastrella; Histiastrella], Spongodiscus Ehrenberg 1854 [Spongodisculus], Spongotrochus Haeckel 1861 [Spongotrochiscus].
          • Euchitoniidae Stöhr 1880 emend. Suzuki 2021: Amphicraspedum Haeckel 1882 [Amphicraspedon; Aphirrhopella], Dictyocoryne Ehrenberg 1861 [Dictyocorynula; Dictyastrum; Dictyastrella; Euchitonia; ?Hymenastromma; Rhopalodictya; Rhopalodictyum; Styla; Pteractis], Hexinastrum Haeckel 1882 [Hexalastromma; Pentalastromma; Pentinastrum], Ommatocampe Haeckel 1861 [Ommatocampium; Amphymenium], Tessarastrum Haeckel 1887 [Tessarastrella; Hagiastromma; Tessarostromma], Tricanastrum Haeckel 1879 [Dicranaster; Dicranastrum; Myelastrella; Myelastrum; Spongomyelastrum; Myelastromma; Pentophiastromma; Spongodicranastrum; Spongohagiastrum; Spongopentophiastrum; Spongostaurina; Tetracranastrum; Triastrum], Trigonastrum Haeckel 1887 [Trigonastrella; Rhopalastromma; Chitonastromma; Amphicraspedina; Amphirrhopoma; Dictyastromma; Monaxonium; Trigonastromma].
          • Spongobrachiidae Haeckel 1882 emend. Suzuki 2021: Spongasteriscus Haeckel 1862 [Spongasterisculus; Dictyocorynium], Spongastromma Haeckel 1887, Spongobrachium Haeckel 1882.
        • Cladococcoidea Haeckel 1862 stat. nov. Suzuki et al. 2021
          • Ethmosphaeridae Haeckel 1862: Cyrtidosphaera Haeckel 1861, Ethmosphaera Haeckel 1861, [Ethmosphaerella; Monosphaera], Haplosphaera Hollande & Enjumet 1960, Heliosphaera Haeckel 1861 [Heliosphaerella], Liosphaera Haeckel 1887 [Melitomma, Craspedomma].
          • Cladococcidae Haeckel 1862: Arachnosphaera Haeckel 1861 [Arachnosphaerella], Arachnospongus Mast 1910, Cladococcus Müller 1856 [Cladococcalis; Anomalacantha; Cladococcodes; Cladococcurus; Porococcus], Diplosphaera Haeckel 1861 [Diplosphaeromma; Astrosphaera; Diplosphaerella; Leptosphaera; Leptosphaerella; Astrosphaerella; Astrospongus; Drymosphaeromma; Leptosphaeromma], Haeckeliella Hollande & Enjumet 1960, Lychnosphaera Haeckel 1882 [Rhizoplegmidium, Rhizospongus, Thalassoplegma].
        • Trematodiscoidea Haeckel 1862 emend. Suzuki 2021 → Trematodiscidae Haeckel 1862 emend. Suzuki 2021: Flustrella Ehrenberg 1839 [Centrospira; Discospirella; Trematodiscus; Perichlamydium; Perispirella; Stylochlamyum], Staurospira Haeckel 1887 [Staurodictyon; Tholodiscus; Xiphospira], Stylodictya Ehrenberg 1846 [Stylodictyon; Stylochlamydium; Stylochlamys; Stylospongia; Stylocyclia; Stylospira], Tripodictya Haeckel 1882 [Xiphodictyon].
        • Haliommoidea Ehrenberg 1846
          • Actinommidae Haeckel 1862 emend. Suzuki 2021: Actinomma Haeckel 1861 [Actinommetta; Haliommura; ?Rhaphidococcus; Riedelipyle; Dreyerella; Drymyomma; Cromyomma; Echinommura; Heliosomura; Spheropyle], Cromyechinus Haeckel 1882 [Cromyodrymus], Rhaphidocapsa Haeckel 1887, ?Sphaeractis Brandt in Wetzel 1936, ?Staurocaryum Haeckel 1882, Stomatosphaera Dreyer 1889, ?Stuermeria Deflandre 1964.
          • Haliommidae Ehrenberg 1846 emend. Suzuki 2021: Cromyosphaera Haeckel 1882 [Cromyommetta; Cromyommura], Haliomma Ehrenberg 1839 [Actinommilla; Cenosphaera; Circosphaera; Thecosphaerella], Haliommantha Haeckel 1887, Hexacontella Haeckel 1887, Melittosphaera Haeckel 1882 [Conoactinomma], Pseudostaurosphaera Krasheninnikov 1960 [Pseudostaurolonche].
          • Heliodiscidae Haeckel 1882 sensu De Wever et al. 2001: Actinommura Haeckel 1887 [?Excentrosphaerella], Excentrococcus Dumitrica 1978, Excentrodiscus Hollande & Enjumet 1960, Heliodiscus Haeckel 1862 [Heliodiscilla, Heliocladus, Heliodiscetta, Heliodrymus, Heliodendrum, Heliosestilla], Helioferrusa Dumitrica 2019, Phaenicosphaera Haeckel 1887 [Dreyeropyle].
        • Lithelioidea Haeckel 1862 sensu Matsuzaki et al. 2015
          • ?†Conocaryommidae Lipman 1969: Conocaryomma Lipman 1969 [?Conocromyomma].
          • Litheliidae Haeckel 1862 emend. Suzuki 2021: Lithelius Haeckel 1861 [Lithospira; ?Azerbaidjanicus], Middourium Kozlova 1999 [Monobrachium], Spiremaria Kozlova 1960 [Spiromultitunica], Spongocyclia Haeckel 1862 [?Lithocarpium; Ommatodiscinus; Ommatodiscus; Ommatodisculus].
          • †Phaseliformidae Pessagno 1972: Phaseliforma Pessagno 1972.
          • †Pyramispongiidae Kozur & Mostler 1978 sensu O'Dogherty 1994: Pyramispongia Pessagno 1973 [Nodotetraedra].
          • Sponguridae Haeckel 1862: Ommatogramma Ehrenberg 1861 [Spongurus; Spongocorisca; ?Spongurantha; Spongurella].
        • Spongopyloidea Dreyer 1889 emend. Suzuki 2021
          • Spongopylidae Dreyer 1889 emend. Suzuki 2021: Schizodiscus Dogiel in Dogiel & Reshetnyak 1952, Spongobrachiopyle Kozur & Mostler 1978, Spongopyle Dreyer 1889 [Spongopylarium], Spongospira Stöhr 1880.
          • Cristallosphaeridae Popofsky 1912: Calcaromma Thomson 1877 [Cristallosphaera], Enalomelon Sugiyama 1992.
          • ?Prunopylidae Poche 1913: Prunopyle Dreyer 1889, Spongopylidium Dreyer 1889.
        • Phorticioidea Haeckel 1882 stat. nov. Suzuki et al. 2021 [Larnacillilae De Wever et al. 2001]
          • Amphitholidae Haeckel 1887 stat. nov. sensu De Wever et al. 2001
          • Circodiscidae Dumitrica 1989 stat. nov. Suzuki et al. 2021
          • Cryptolarnaciidae Dumitrica 1989 stat. nov. Suzuki et al. 2021
          • Histiastridae Dumitrica 1989 stat. nov. Suzuki et al. 2021
          • Phorticiidae Haeckel 1882 sensu Dumitrica 1989
        • Larcospiroidea Haeckel 1887 stat. nov. sensu Dumitrica 1989
          • Dipylissidae Dumitrica 1989 stat. nov. Suzuki et al. 2021
          • Larcospiridae Haeckel 1887 stat. nov. Suzuki et al. 2021
          • Palaeotetrapylidae Dumitrica 1989 stat. nov. Suzuki et al. 2021
          • Pylodiscidae Haeckel 1887 sensu Dumitrica 1989
          • Zonariidae Haeckel 1887 sensu Dumitrica 1989
        • Pseudoaulophacoidea Riedel 1967 sensu De Wever et al. 2001
          • Patulibracchiidae Pessagno 1971 sensu De Wever et al. 2001
          • Pseudoaulophacidae Riedel 1967 sensu De Wever et al. 2001
          • Suttoniidae Schaaf 1976 sensu Dumitrica 2019
        • Stylosphaeroidea Haeckel 1887 sensu Dumitrica 1984
          • Entapiidae Dumitrica in De Wever, Dumitrica, Caulet Nigrini & Caridroit 2001
          • Stylatractidae Schröder 1909 emend. Suzuki 2021
          • Tubosphaeridae Suzuki 2021
      • Nassellaria Ehrenberg 1875 emend. Haeckel 1887. Artostrobus, Eucyrtidium, Lithomelissa, Pterocanium, Pterocorys.
      • Collodaria Haeckel 1887. Acrosphaera, Collosphaera, Collozoum, Sphaerozoum, Rhaphidozoum, Siphonsphaera, Thalassicolla.
      • Orodaria Nakamura & Suzuki 2020[159]
        • Oroscenidae Haeckel 1887 emend. Nakamura & Suzuki 2021 [Orosphaerida Haeckel 1887; Orosphaeridae Haecker 1908]. Oroscena, Orostaurus, Oropelex. Nomina dubia genera: Orosphaera, Orodictyum, Orona, Oronium.
        • Thalassothamnidae Haecker 1906 emend. Nakamura & Suzuki 2021 [Cytocladidae Schröder, 1908]. Thalassothamnus, Cytocladus. Nomen dubium genus: Conostylus.
      • †Entactinaria Kozur & Mostler 1982 sensu Nakamura & Suzuki 2020[v]
        • †Entactiniidae Riedel 1967. Afanasievella, Apophysiactinia, Borisella, Callela, Cancellosphaera, Costaentactinia, Cyclocarpus nomen dubium, Duodecimentactinia, Langsonsphaera,[160] Entactinosphaera, Gracilentactinia, Holdisphaera, Involutentactinia, Kashiwara, Magnentactinia, Magnisphaera, Microporosa, Moskovistella, Multientactinia, Munzuwonella, Paratriposphaera, Perforentactinia, Plenoentactinia, Plenosphaera, Pluristratoentactinia, Polyedroentactinia, Provisocyntra, Radiobisphaera, Retentactinia, Retisphaera, Sinosphaera, Spongentactinia, Stigmosphaerostylus, Trilonche, Uberinterna, Wuyia.[161]
        • †Tetrentactiniidae Kozur & Mostler 1979. Multisphaera.
        • †Triassothamnidae Takahashi, Maekawa & Dumitrica 2022. Triassothamnus.[160]
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Excavates

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Perspective

Malawimonadida

Metamonada

The metamonads were first described by Pierre-Paul Grassé in the first volume of Traité de Zoologie, published in 1952, as the superorder Metamonadina or Anaxostylaria. They were originally composed of zooflagellates with four or more flagella, known as polymastigotes and hypermastigotes (e.g., Trichomonas, Oxymonas).[163] These were later split into other groups such as the Parabasalia and Preaxostyla (over 260 and 140 species respectively),[164][165] regarded as individual phyla. After rRNA phylogenetic analyses demonstrated their relatedness, the concept of metamonads was reintroduced by Cavalier-Smith as a monophyletic phylum Metamonada that includes both groups, as well as the anaerobic Fornicata (e.g., the free-living Carpediemonas and the parasitic Giardia;[166][167] around 140 species).[164] In this system, Preaxostyla is known as subphylum Anaeromonada, and Fornicata and Parabasalia are two infraphyla that belong to the subphylum Trichozoa.[7] Still, some taxonomists retain the Parabasalia at a phylum level.[168]

The classifications of Preaxostyla and Fornicata were last revised separately in the 2017 Handbook of the protists[169][170][171] and the higher classification of Metamonada was revised by Cavalier-Smith in 2021.[7] The classification of Parabasalia was completely updated in 2024.[168] Two new smaller clades of metamonads have been described in addition: anaeramoebae[172] and the 'BaSk' clade, containing barthelonids and skoliomonads.[173] However, Parabasalia was treated as a separate phylum by the 2024 revision, instead of a member of phylum Metamonada.[168] For consistency, the classification listed below maintains Parabasalia as part of the phylum Metamonada.

Phylum Metamonada Grassé 1952 emend. Cavalier-Smith 2003

  • Family Anaeramoebidae Táborský, Pánek & Čepička 2017, sole genus Anaeramoeba.
  • 'BaSk' clade, genera Barthelona, Skoliomonas.
  • Subphylum Preaxostyla (=Anaeromonada Cavalier-Smith 1996 emend. 2003), class Anaeromonadea Cavalier-Smith 1996 emend. 2003
    • Order Trimastigida Cavalier-Smith 2003 emend. 2021, family Trimastigidae Kent 1880 emend. Zhang et al. 2015, sole genus Trimastix.
    • Order Paratrimastigida Cavalier-Smith 2021, family Paratrimastigidae Zhang, Taborsky, Silberman, Panek, Čepička & Simpson 2015, sole genus Paratrimastix.
    • Order Oxymonadida Grassé 1952. Genus not assigned to any family: Opisthomitus.[w]
      • Family Oxymonadidae, genera Barroella, Oxymonas, Microrhopalodina, Sauromonas, †Oxymonites, †Microrhopalodites, †Sauromonites.
      • Family Polymastigidae, genera Blattamonas, Brachymonas, Monocercomonoides, Paranotila, Polymastix, Tubulimonoides.
      • Family Pyrsonymphidae, genera Dinenympha, Pyrsonympha, †Dinenymphites, †Pyrsonymphites.
      • Family Saccinobaculidae, genera Notila, Saccinobaculus.
      • Family Streblomastigidae, sole genus Streblomastix.
  • Subphylum Trichozoa(P?) Cavalier-Smith 1996 emend. 2003
    • Infraphylum Fornicata Simpson 2003[x]
      • "Carpediemonas-like organisms"(P) Kolisko et al. 2010 (class Carpomonadea Cavalier-Smith 2013), genera Aduncisulcus, Carpediemonas, Ergobibamus, Hicanonectes, Kipferlia.
        • Order Dysnectida Cavalier-Smith 2013, family Dysnectidae Cavalier-Smith 2013, sole genus Dysnectes.
        • Order Chilomastigida Cavalier-Smith 2013, family Chilomastigidae Cavalier-Smith 2013, sole genus Chilomastix.
      • Class Eopharyngea Cavalier-Smith 1993
    • Infraphylum Parabasalia Honigberg 1973. Genera incertae sedis: Chilomitus, Rhizonympha, Tricercomitus, Trichocovina.
      • Class Hypotrichomonadea Čepička, Hampl & Kulda 2010, order Hypotrichomonadida Čepička, Hampl & Kulda 2010, family Hypotrichomonadidae Honigberg 1963 emend. Čepička, Hampl & Kulda 2010, genera Hypotrichomonas, Trichomitus.
      • Class Pimpavickea Boscaro & Keeling 2024, order Pimpavickida Céza & Čepička 2022, family Pimpavickidae Céza & Čepička 2022, genera Pimpavicka, Alexandriella.
      • Class Trichomonadea Kirby 1947 sensu Čepička, Hampl & Kulda 2010
        • Order Honigbergiellida Čepička, Hampl & Kulda 2010 emend. Boscaro & Keeling 2024
          • Family Hexamastigidae Čepička, Hampl & Kulda 2010 emend. Boscaro & Keeling 2024, sole genus Hexamastix.
          • Family Honigbergiellidae Čepička, Hampl & Kulda 2010, genera Honigbergiella, Ditrichomonoides, Honigbergiellopsis, Monotrichomonas.
          • Family Cthulhuidae Boscaro & Keeling, genera Cthulhu, Cthylla, Nyarlathotep.
          • Family Tetratrichomastigidae Boscaro & Keeling 2024, sole genus Tetratrichomastix.
        • Order Trichomonadida Kirby 1947 sensu Čepička, Hampl & Kuda 2010. Genera incertae sedis: Pseudotrichomonas, †Paleotrichomones.[177]
          • Family Lacusteriidae Céza & Čepička 2022, sole genus Lacusteria.
          • Family Trichomonadidae Chalmers & Pekkola 1918 sensu Hampl, Vrlík, Čepička, Pecka, Kulda & Tachezy 2006, genera Trichomonas, Cochlosoma, Cyathosoma, Pentatrichomonas, Pentatrichomonoides, Pseudotrypanosoma, Ptychostoma, Tetratrichomonas, Trichomitopsis, Trichomonoides.
      • Clade "Tla"
        • Class Lophomonadea Boscaro & Keeling 2024, order Lophomonadida Light 1927 emend. Cavalier-Smith 2013, family Lophomonadidae Kent 1880 emend. Cavalier-Smith 2013, genera Lophomonas, Eulophomonas, Prolophomonas.
        • Class Trichonymphea Poche 1913, order Trichonymphida Poche 1913
          • †Family Burmanymphidae Poinar 2009, sole genus Burmanympha.[177]
          • Family Barbulanymphidae Boscaro & Keeling 2024, genera Barbulanympha, Rhynchonympha, Urinympha.
          • Family Hoplonymphidae Light 1926 emend. Boscaro & Keeling 2024, sole genus Hoplonympha.
          • Family Retractinymphidae Radek & Brune 2023, sole genus Retractinympha.
          • Family Spirotrichosomidae Hollande & Carruette-Valentin 1971, genera Spriotrichosoma, Apospironympha, Bispironympha, Colospironympha, Heliconympha, Leptospironympha, Macrospironympha.
          • Family Staurojoeninidae Grassi 1917, genera Staurojoenina, Idionympha.
          • Family Teranymphidae Koidzumi 1921, genera Tetranympha, Eucomonympha, Pseudotrichonympha.
          • Family Trichonymphidae Kent 1880, sole genus Trichonympha.
      • Clade "Cadamassta"
        • Class Cristamonadea Čepička, Hampl & Kulda 2010. Genera incertae sedis: Cyclojoenia, Joenia, Joenina, Joenoides, Joenopsis, Pachyjoenia, Parajoenopsis, Placojoenia, Projoenia.
          • Order Calonymphida Boscaro & Keeling 2024
            • Family Calonymphidae Grassi 1911 sensu Gile 2011, genera Calonympha, Criconympha, Daimonympha, Diplonympha, Gyronympha, Metastephanonympha, Prosnyderella, Snyderella, Stephanonympha.
            • Family Deltotrichonymphidae Brugerolle & Lee 2000, genera Deltotrichonympha, Koruga.
            • Family Mixotrichidae Boscaro & Keeling 2024, sole genus Mixotricha.
          • Order Devescovinida Boscaro & Keeling 2024, family Devescovinidae Doflein 1911 emend. Boscaro & Keeling 2024, genera Devescovina, Achemon, Astronympha, Bullanympha, Caduceia, Coronympha, Evemonia, Foaina, Hyperdevescovina, Kirbynia, Kofoidia, Macrotrichomonas, Macrotrichomonoides, Metadevescovina, Parajoenia, Polymastigoides, Pseudodevescovina, Runanympha, †Devescovites.[177]
          • Order Gigantomonadida Boscaro & Keeling 2024, family Gigantomonadidae Boscaro & Keeling 2024, sole genus Gigantomonas.
        • Class Dientamoebea Boscaro & Keeling 2024, order Dientamoebida Boscaro & Keeling 2024, family Dientamoebidae Grassé 1953, genera Dientamoeba, Histomonas, Parahistomonas, Protrichomonas.
        • Class Monocercomonadea Boscaro & Keeling 2024, order Monocercomonadida Boscaro & Keeling 2024, family Monocercomonadidae Kirby 1944, sole genus Monocercomonas.
        • Class Simplicimonadea Boscaro & Keeling 2024, order Simplicimonadida Boscaro & Keeling 2024, family Simplicimonadidae Čepička, Hampl & Kulda 2010, sole genus Simplicimonas.
        • Class Tritrichomonadea Čepička, Hampl & Kulda 2010 emend. Boscaro & Keeling 2024, order Tritrichomonadida Čepička, Hampl & Kulda 2010 emend. Boscaro & Keeling, family Tritrichomonadidae Čepička, Hampl & Kulda 2010, sole genus Tritrichomonas.
        • Class Spirotrichonymphea Grassé 1952 sensu Čepička, Hampl & Kulda 2010
          • Order Cononymphida Boscaro & Keeling 2024, family Cononymphidae Boscaro & Keeling 2024, sole genus Cononympha.
          • Order Holomastigotoidida Boscaro & Keeling 2024, family Holomastigotoididae Grassi & Foà 1911, genera Holomastigotoides, Rostronympha.
          • Order Spirotrichonymphida Grassé 1952
            • Family Brugerollinidae Boscaro & Keeling 2024, genera Brugerollina, Pseudospironympha.
            • Family Fraterculidae Boscaro & Keeling 2024, genera Fraterculus, Cuppa.
            • Family Holomastigotidae Janicki 1915, genera Holomastigotes, Spiromastigotes, Spirotrichonymphella, Uteronympha.
            • Family Spirotrichonymphidae Grassi 1917, genera Spirotrichonympha, Microjoenia, Micromastigotes, Nanospironympha, Spironympha, Torquenympha.

Discoba

Jakobida

Tsukubamonadida Yabuki et al. 2011

Heterolobosea

Pharyngomonadida

Tetramitia
Selenaionea

Neovahlkampfiida

Selenaionida

Eutetramitea

Naegleriida

Percolomonadida

Pseudociliatida

Creneida

Lyromonadida

Cladogram of heterolobosean orders based on combined phylogenomic and 18S rRNA analyses published in 2025.[181]

The phylum Heterolobosea contains around 170 species of amoebae, flagellates, and amoeboflagellates. It was initially established to unite two historically well-known amoeboid orders, Schizopyrenida (such as Naegleria fowleri, a human pathogen) and Acrasida (slime molds). Later, as more flagellates joined this grouping, the usage of Heterolobosea was split between two meanings: the more common usage applies to the entire clade,[6] while the usage by Cavalier-Smith and collaborators was restricted to a paraphyletic class of "traditional" heteroloboseans, with the name Percolozoa used for the phylum instead.[182][183] The first comprehensive phylogenomic study of Heterolobosea was published in 2025 by Tomáš Pánek and coauthors, resulting in its modern classification.[181]

Phylum Heterolobosea Page & Blanton 1985 sensu Hanousková et al. 2019 (=Percolozoa Cavalier-Smith 1991)

  • Subphylum Pharyngomonada Cavalier-Smith 2008
    • Class Pharyngomonadea Cavalier-Smith in Cavalier-Smith & Nikolaev 2008
      • Order Pharyngomonadida Cavalier-Smith in Cavalier-Smith & Nikolaev 2008
        • Family Pharyngomonadidae Cavalier-Smith in Cavalier-Smith & Nikolaev 2008, sole genus Pharyngomonas.
  • Subphylum Tetramitia Cavalier-Smith 1993
    • ?Family Orodruinidae Shɨshkin 2021,[y] genera Gruberella, Orodruina, Stachyamoeba.
    • Class Selenaionea Pánek & Čepička 2025
      • Order Neovahlkampfiida Cavalier-Smith 2022 emend. Pánek et al. 2025
      • Order Selenaionida Hanousková et al. 2018
        • Family Selenaionidae Hanousková et al. 2018, sole genus Selenaion.
    • Class Eutetramitea Pánek & Čepička 2025. Genera not assigned to any order and family: Euplaesiobystra, Fumarolamoeba, Heteramoeba, Parafumarolamoeba, Paravahlkampfia, Oramoeba, Vrihiamoeba.
      • Order Naegleriida Pánek & Čepička 2025 (=Heterolobosea sensu Cavalier-Smith 2022). Genera not assigned to any family: Aurem, Marinamoeba.
      • Order Percolomonadida Cavalier-Smith 1993
        • Family Barbeliidae Arndt 2023, genera Barbelia, Nonamonas.
        • Family Lulaidae Hohlfeld & Arndt 2023, sole genus Lula.
        • Family Percolomonadidae Cavalier-Smith 2003, sole genus Percolomonas.
      • Order Pseudociliatida Corliss & Lipscomb 1982
      • Order Creneida Cavalier-Smith 2022
        • Family Creneidae Pánek et al. 2014, sole genus Creneis.
      • Order Lyromonadida Cavalier-Smith 1993

Euglenozoa

The phylum Euglenozoa is home to at least 2,000 described species of single-celled flagellates of very dissimilar lifestyles. It was originally proposed to group the euglenids (such as the photosynthetic Euglena) and the kinetoplastids (like the pathogenic Trypanosoma), usually studied separatedly; eventually it included diplonemids and symbiontids as well.[184][185][186] Due to its share of photosynthetic species, traditionally regarded as algae, the phylum is also known as Euglenophyta by phycologists, and euglenids in particular were often studied as algae. Euglenids and kinetoplastids are the most diverse in terms of described species, although diplonemids may compose over 67,000 potential species.[186] The classification of euglenozoans was summarized by Cavalier-Smith in 2016.[185] A more phylogenetically precise revision of their classification was published in 2021 by Alexei Kostygov and coauthors,[186] with newer clades and genera described in the following years, particularly of euglenids. Still, one group of euglenids, the paraphyletic "ploeotids",[187] remains unresolved in the current classification, spread out across multiple clades.[188][189][190]

Phylum Euglenozoa Cavalier-Smith 1981 emend. Simpson 1997

  • Class Euglenida Bütschli 1884 emend. Simpson 1997 (=Euglenoida Bütschli 1884 emend. Cavalier-Smith 1993). Genera not assigned to any lower clade: †Moyeria,[191] Entosiphon, Gaulosia. Genera incertae sedis: Atraktomonas*, Calycimonas*, Dolium*, Dylakosoma*, Tropidoscyphus*, Michajlowastasia*, Parastasiella*, Dinemula*, Paradinemula*, Mononema*, Ovicola*, Embryocola*, Copromonas*.
    • Order Petalomonadida Cavalier-Smith 1993, family Scytomonadidae Stein 1878 (=Petalomonadidae Bütschli 1884), genera Biundula, Notosolenus, Pentamonas*, Petalomonas, Scytomonas, Sphenomonas.
    • Alistosa Lax & Simpson 2021, genera Ploeotia, Serpenomonas, Keelungia, Lentomonas, Decastava.
    • Karavia Lax, Cho & Keeling 2023, genera Hemiolia, Liburna.
    • Olkaspira Lax & Simpson 2021. Genera not assigned to any lower clade: Chelandium, Olkasia.
      • Spirocuta Cavalier-Smith, Chao & Vickerman 2016 (=Helicales Perschke et al. 2017)
        • Anisonemia Cavalier-Smith 2016
          • Order Anisonemida Cavalier-Smith 2016, family Anisonemidae Kent 1880 emend. Cavalier-Smith 2016, genera Anisonema, Dinema (=Dinematomonas), Heteronema.
          • Order Natomonadida Cavalier-Smith 2016
            • Family Neometanemidae Cavalier-Smith 2016, sole genus Neometanema.
            • Aphagea Cavalier-Smith 1993 emend. Busse & Preisfeld 2003 (=Rhabdomonadina Leedale 1967 emend. Cavalier-Smith 1993)
              • Family Astasiidae Kent 1884, genera Astasia, Gyropaigne, Menoidium, Parmidium, Rhabdomonas.
              • Family Distigmidae Hollande 1942, sole genus Distigma.
        • Order Peranemida (Peranematales) Cavalier-Smith 1993, family Peranemidae (Peranemataceae) Bütschli 1884, genera Chasmostoma, Jenningsia (=Peranemopsis),[192] Peranema, Teloprocta, Urceolus (=Urceolopsis, Phialonema).[193][194]
        • Euglenophyceae Schoenichen in Eyfurth & Shoenichen 1925, emend. Marin & Melkonian 2003 (=Euglenea Butschli 1884, emend. Busse & Preisfeld 2002). Genera incertae sedis: Ascoglena*, Euglenamorpha*, Euglenopsis*, Glenoclosterium*, Hegneria*, Klebsina*, Euglenocapsa*.
  • Class Symbiontida Yubuki, Edgcomb, Bernhard & Leander 2009 (=Postgaardea Cavalier-Smith, 1998), genera Bihospites, Calkinsia, Postgaardi.
  • Class Diplonemea Cavalier-Smith 1993, order Diplonemida Cavalier-Smith 1993
    • Family Diplonemidae Cavalier-Smith 1993 emend. Adl et al. 2019, genera Diplonema, Rhynchopus, Lacrimia, Sulcionema, Flectonema.
    • Family Hemistasiidae Cavalier-Smith 2016 emend. Adl et al. 2019, genera Hemistasia, Namystinia, Artemidia.
    • Family Eupelagonemidae Okamoto & Keeling 2018, sole genus Eupelagonema.
  • Class Kinetoplastea Honigberg 1963 emend. Vickerman 1976. Genera incertae sedis: Bordnamonas*, Cephalothamnium*, Desmomonas*, Jarrellia*, Lamellasoma*.
    • Subclass Prokinetoplastia Vickerman in Moreira et al. 2004, order Prokinetoplastida Vickerman in Moreira et al. 2004
      • Family Ichthyobodonidae Isaksen et al. 2007, sole genus Ichthyobodo.
      • Family Perkinselidae Kostygov in Kostygov et al. 2021, sole genus Perkinsela.
    • Subclass Metakinetoplastia Vickerman in Moreira et al. 2004
      • Family Allobodonidae Goodwin, Lee, Kugrens & Simpson 2018,[195] genera Allobodo, Novijibodo.[196]
      • Order Neobodonida Vickerman in Moreira et al. 2004
        • Family Neobodonidae(P?) Cavalier-Smith 2016, genera Actuariola, Avlakibodo,[197] Azumiobodo, Cruzella, Cryptaulaxella, Klosteria, Neobodo, Phanerobia, Rhynchobodo.
        • Family Rhynchomonadidae Cavalier-Smith 2016, genera Dimastigella, Rhynchomonas.
      • Order Parabodonida Vickerman in Moreira et al. 2004
        • Family Cryptobiidae Poche 1911 emend. Kostygov in Kostygov et al. 2021, genera Cryptobia, Parabodo.
        • Family Trypanoplasmatidae Hartmann & Chagas 1910 emend. Kostygov in Kostygov et al. 2021, genera Procryptobia, Trypanoplasma.
      • Order Eubodonida Vickerman in Moreira et al. 2004, family Bodonidae Bütschli 1883, sole genus Bodo.
      • Order Trypanosomatida Kent 1880 emend. Vickerman in Moreira et al. 2004, family Trypanosomatidae Dolfein 1901. Genera not assigned to subfamilies: Jaenimonas, Vickermania, Sergeia, Wallacemonas. Genera incertae sedis: Cercoplasma, Malacozoomonas, Nematodomonas, Rhynchoidomonas, †Paleoleishmania G.Poinar & R.Poinar 2004.[198]
        • Subfamily Trypanosomatinae Doflein 1901, sole genus Trypanosoma.
        • Subfamily Leishmaniinae Maslov & Lukeš 2012
          • Infrafamily Crithidiatae Kostygov & Yurchenko 2017, genera Leptomonas, Crithidia, Lotmaria.
          • Infrafamily Leishmaniatae Maslov & Lukeš 2012, genera Leishmania, Porcisia, Endotrypanum, Novymonas Zelonia, Borovskyia.
        • Subfamily Herpetomonadinae Alexeieff 1911 stat. nov., emend. Kostygov & Yurchenko in Kostygov et al. 2021 (=Phytomonadinae Yurchenko, Kostygov, Votýpka & Lukeš 2015), genera Herpetomonas, Lafontella, Phytomonas.
        • Subfamily Strigomonadinae Votýpka, Yurchenko, Kostygov & Lukeš 2014, genera Angomonas, Strigomonas, Kentomonas.
        • Subfamily Blastocrithidiinae Votýpka, Yurchenko & Lukeš 2021, genera Blastocrithidia, Obscuromonas.
        • Subfamily Blechomonadinae Votýpka & Suková 2013, sole genus Blechomonas.
        • Subfamily Paratrypanosomatinae Votýpka & Lukeš 2013, sole genus Paratrypanosoma.
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Minor clades

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Perspective

Ancyromonadida

  • Divimonas Barkhouse, Eglit, Weston & Simpson 2025[200]
  • Ancyromonadidae Cavalier-Smith 2008. Ancyromonas, Nutomonas.
  • Planomonadidae Cavalier-Smith 1993. Fabomonas, Planomonas.

CRuMs

Hemimastigophora

Provora

Telonemia

The phylum Telonemia (telonemids) contains a few species of flagellates found in ocean and fresh waters worldwide. It was originally proposed in 2006 for Telonema, a genus of previously uncertain affinity.[205] Under the Cavalier-Smith system, telonemids were initially classified as a class of Cryptista,[66] but later analyses consistently recovered it as a separate group.[14] Until 2019, only two species had been formally described, each belonging to a separate genus,[206] although environmental DNA sequencing suggests there are many more species not yet described.[207] In 2022, five additional species were described along with a third new genus, bringing the total number of species to seven.[208]

Phylum Telonemia Shalchian-Tabrizi 2006, class Telonemea Cavalier-Smith 1993, order Telonemida Cavalier-Smith 1993, family Telonemidae Cavalier-Smith 1993, genera Arpakorses, Lateronema, Telonema.

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Protists of uncertain affiliation

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Perspective

Acinetactis, Actinastrum, Actinocoma, Actinolophus, Adinomonas, Aletium, Amphimonas, Amylophagus, Aphelidiopsis, Asterocaelum, Asthmatos, Aurospora, Barbetia, Berkeleyaesol,[209] Belaria, Belonocystis, Bertarellia, Bertramia, Bodopsis, Boekelovia, Branchipocola, Camptoptyche, Chalarodora, Cibdelia, Cichkovia, Cinetidomyxa, Cingula, Cladomonas, Clathrella, Codonoeca, Coelosporidium,[z] Copromonas, Cyanomastix, Cyclomonas, Cytamoeba, Dallingeria, Dictyomyxa, Dimastigamoeba, Dinemula, Dinoasteromonas, Diplocalium, Diplomita, Diplophysalis, Diploselmis, Dobellina, Ducelleria, Ectobiella, Elaeorhanis, Embryocola, Endamoeba, Endemosarca, Endobiella, Endomonas, Endospora, Enteromyxa, Eperythrocytozoon, Errera, Fromentella, Gweamonas,[210] Gymnococcus, Gymnophrydium, Haematotractidium, Hartmannina, Heliobodo, Heliomonas, Hermisenella, Heterogromia, Hillea, Hyalodaktylethra, Immanoplasma, Isoselmis, Janickina, Kamera, Labyrinthomyxa Dubosq 1921,[aa] Lagenidiopsis, Liegeosia, Luffisphaera,[ab] Lymphocytozoon, Lymphosporidium, Macappella, Magosphaera, Malpighiella, Martineziella, Megamoebomyxa, Meringosphaera, Microcometes, Monochrysis, Monodus, Mononema, Myrmicisporidium, Naupliicola, Nephrodinium, Neurosporidium, Orbulinella, Ovicola, Palisporomonas, Pansporella, Paradinemula, Parakaryon,[ac] Paraluffisphaera, Paramonas, Paraplasma, Parastasia, Parastasiella, Peliainia, Peltomonas, Petasaria, Phagodinium, Phanerobia, Phloxamoeba, Phyllomitus, Phyllomonas, Physcosporidium, Pleuophrys, Pleuromastix, Protenterospora, Protomonas, Pseudoactiniscus, Pseudosporopsis, Rhizomonas, Rhynchodinium, Rigidomastix, Schewiakoffia, Sergentella, Serpentoplasma, Sphaerasuctans, Spongastericus, Spongocyclia, Stephanomonas, Strobilomonas, Tetradimorpha, Thaulirens, Topsentella, Toshiba, Trichonema, Urbanella.[6]

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See also

Notes

  1. The first eukaryotes were “neither plants, animals, nor fungi”, hence as defined, the Protista would include the earliest common ancestor of all eukaryotes.
  2. In the original 4-kingdom model proposed in 1959, Protista included all unicellular microorganisms such as bacteria.
  3. The position of the genera Microcorycia, Parmulina, Penardochlamys and Zonomyxa, which were listed in 2002 under family Microcoryciidae, is not clear. They are placed here by morphological characters but this needs to be supported by molecular data.[6]
  4. This genus grouped with Dermamoebidae in a phylogenetic anaysis that used a limited number of taxa.[32] In the recent taxonomy it was listed as a separate clade until its position was better resolved.[6] A 2023 analysis placed it, once again, related to Dermamoebida in a clade with Microglomus.[33]
  5. The two genera of Stygamoebidae, Stygamoeba and Vermistella, group together in some phylogenetic analyses, but usually appear separated. In 2019 they were considered only a potential branch of Discosea, unless proven otherwise.[6] A 2023 analysis places both genera in Flabellinia, closely related to Thecamoebida.[33]
  6. The orders Thecamoebida, Dactylopodida, Acanthopodida, Himatismenida and Pellitida were initially divided into families, but phylogenetic analyses haven't supported the monophyly of each family. For this reason, in recent classifications there is no family rank division within it.[25][6]
  7. The genus Pessonella could be a synonym of Vannella.[6]
  8. The 2019 revision by the ISOP ignores the grouping of some variosean genera into higher rank clades (orders and families) proposed in older studies, due to the weakly supported SSU rRNA phylogenetic analyses.[6]
  9. The picobiliphytes are phylogenetically closer to Rhodophyta and Rhodelphis than to other groups, and are therefore considered part of Archaeplastida.[44]
  10. The 2019 revision by the ISOP inaccurately stated that goniomonads are classified as "Cyathomonadacea Pringsheim 1944".[6] The name Cyathomonadacea does not exist; Pringsheim only described the family Cyathomonadaceae based on the genus Cyathomonas, and later the order Cyathomonadales and class Cyathomonadea were described, each by different authors. However, as pointed out in 1993 by Gianfranco Novarino and Ian Lucas, this genus was based on a species that actually belonged to Goniomonas; since then, taxonomists have prioritized higher taxa named after it (Goniomonadaceae, Goniomonadales/adida, Goniomonadea) instead.[67]
  11. The genus Tetragonidium is known only from one written diagnosis and some illustrations, and its affinities with cryptomonads are very uncertain.[67]
  12. Within the order Anoecida, the four families Anoecaceae, Caecitellaceae, Cafeteriaceae and Symbiomonadaceae were included by T. Cavalier-Smith in 2006. In a 2013 revision, he simplified the classification by transferring Symbiomonadaceae and Anoecaceae to Cafeteriaceae.[79] However, both of those families remain accepted by the scientific community as independent from Cafeteriaceae as of 2020.[77]
  13. Cavalier-Smith hypothesized in 2013 a sister relationship between actinophryids and raphidophytes, with both groups placed under the taxon Raphidomonadea. The molecularly uncharacterized genus Commation was classified as part of this taxon.[79] However, the placement of actinophryids among ochrophytes is still unstable and uncertain,[90][91] and Commation has never been included in classifications since.[citation needed]
  14. O'Kelly placed the genera Antarctosaccion, Chrysomeris, Chrysonephos, Nematochrysis/Chrysowaernella, Phaeosaccion and Rhamnochrysis in the order Chrysomeridales nomen nudum in 1989,[93] and Cavalier-Smith placed it inside the class Chrysomeridophyceae (spelt originally as Chrysomerophycea) in 1995.[94] However, the class was proven to be polyphyletic. As a result, some of its genera (Nematochrysis/Chrysowaerella) were transferred to the class Chrysoparadoxophyceae, while others (Antarctosaccion and Phaeosaccion) were transferred to the class Phaeosacciophyceae. Due to a lack of molecular data, the placement of Chrysomeris, Chrysonephos and Rhamnochrysis remains unknown.[95]
  15. The position of Eustigmatophyceae is still unstable, with different phylogenetic methodologies showing affinities to either of the SI and SII clades.[97][91]
  16. The taxonomic ranks above class level (phylum, subphylum, infraphylum, etc.) are unstable within the Alveolata, due to the many different existing classifications. For example, the phylum-level rank has been traditionally assigned to Perkinsozoa,[111] Dinoflagellata, Chromerida and Apicomplexa, all of which have been classified as phylum Myzozoa,[112] which in turn has been classified inside phylum Miozoa.[84] Treating the four myzozoan groups as separate phyla is generally accepted among phycologists,[113] while at the same time the rejection of higher taxonomic ranks due to being superfluous is prevalent among protistologists.[6]
  17. Apart from morphological descriptions, no molecular analysis has solidly affiliated Rastrimonas with the Perkinsozoa.[114]
  18. The genera Parallobiopsis, Ellobiocystis and Rhizellobiopsis are only provisionally placed among ellobiopsids.[125]
  19. Highly divergent 18S rRNA.[6] Since 2020 member of newly identified major apicomplexan subgroup Marosporida, putting together Aggregata octopiana Frenzel 1885, Merocystis kathae Dakin, 1911 (both Aggregatidae, originally coccidians), Rhytidocystis sp. 1 and Rhytidocystis sp. 2 Janouškovec et al. 2019 (Rhytidocystidae Levine, 1979, originally coccidians, Agamococcidiorida), and Margolisiella islandica Kristmundsson et al. 2011 (closely related to Rhytidocystidae)[130]
  20. This taxon is artificial; many of the families listed in it are not monophyletic and have little support from phylogenetic analyses.[6]
  21. There is only one molecular sequence from this genus, which causes long branches in gene phylogenies, and the node is unresolved.[6]
  22. According to Nakamura & Suzuki, the only extant (i.e. not extinct) orders of Polycystinea are Spumellaria, Nassellaria, Collodaria and Orodaria. Therefore, Entactinaria is considered an exclusively fossil group.[159]
  23. One 18S rRNA phylogenetic analysis suggests that the genus Opisthomitus is affiliated with the family Pyrsonymphidae.[169]
  24. Regarding the classification of Fornicata, there is a disconnect between the Cavalier-Smith system and the system described by different authors in the Handbook of the protists and other studies. The order Retortamonadida was originally defined as two genera: Chilomastix and Retortamonas. As both groups of taxonomists have reported, retortamonads are polyphyletic, because Chilomastix branches distantly from Retortamonas. In 2013, Cavalier-Smith fixed this taxonomic issue by modifying the retortamonads to only include Retortamonas, and describing a separate order Chilomastigida for Chilomastix only, placed among the early branching Carpediemonas-like organisms.[174] However, this has not yet been recognized by later studies from the other authors, as they have continued to report a polyphyletic Retortamonadida.[171][175][176] Because of this inconsistency, Eopharyngea (the clade formed by Retortamonadida plus Diplomonadida) is monophyletic only when considering one version of Retortamonadida, but not the other.
  25. The family Orodruinidae is possibly polyphyletic, and its type genus Orodruina may belong to the order Neovahlkampfiida.[181]
  26. Probably a synonym of the zygomycete fungus Nephridiophaga.[6]
  27. Labyrinthomyxa, a genus of putative labyrinthulomycetes, is placed incertae sedis because of limited data.[211]
  28. May be the same genus as Belonocystis.[6]
  29. It is not yet known whether this organism is a eukaryote or a prokaryote.[212]

References

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