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Taxonomy of Protista

From Wikipedia, the free encyclopedia

A modern non-hierarchical interpretation of the phylogeny of eukaryotes, including protists

A protist (/ˈprtɪst/) 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]

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

Key to symbols used:

(P) = Paraphyletic group.
† = Extinct group.
? = Group of uncertain taxonomic position.
* = Group lacking molecular data.
[ ] = Synonyms and alternative names.

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Transcription

Current classification

This article largely follows the efforts of the scientific community and the International Society of Protistologists to revise the taxonomy of protists in a manner that reflects their phylogeny and evolution, striving away from the use of historical paraphyletic taxa and relying exclusively on clades as the basis of the classification. The reference used for the general arrangement of groups is the publication Revisions to the Classification, Nomenclature and Diversity of Eukaryotes, published in 2019 by researcher Sina Adl and coauthors.[3] Other studies are used for some specific groups of protists that have been revised after 2018, such as slime moulds,[4] glaucophyte algae,[5] arcellinid testate amoebae[6] and planktonic foraminifera.[7]

The following cladogram shows the relationships between the major groups of protists, as well as the kingdoms of animals, plants and fungi, according to the most recent phylogenetic analyses.[8][9][10][11][12]

Eukaryota
Choanozoa (P)
sensu Cavalier-Smith

Amorphea Adl et al. 2012

Obazoa Brown et al. 2013

Cladogram of Obazoa based on several analyses.[11][13] It includes the earliest diverging 'fungi' (microsporidians, aphelids and rozellids), which are only sometimes considered protists under the name of Opisthosporidia.[14]

Incertae sedis Obazoa:

Opisthokonta Cavalier-Smith 1987, emend. Adl et al. 2005

  • Nucletmycea Brown et al. 2009 [Holomycota Liu et al. 2009]
  • Holozoa Lang et al. 2002. Incertae sedis: Tunicaraptor Tikhonenkov, Mikhailov, Hehenberger, Karpov, Prokina, Esaulov, Belyakova, Mazei, Mylnikov, Aleoshin & Keeling 2020,[11] Bicellum Strother & Wellman 2021.[18]
    • Ichthyosporea[d] Cavalier-Smith 1998 [Mesomycetozoea Mendoza et al. 2002]
    • Pluriformea[d] Hehenberger et al. 2017
    • Filasterea Shalchian-Tabrizi et al. 2008. Capsaspora, Ministeria, Pigoraptor, Txikispora.[19]
    • Choanozoa Brunet & King 2017 [Choanozoa Cavalier-Smith et al. 1991 (P)][e]
      • Choanoflagellata Kent 1880–1882 [Craspedomonadina Stein 1878; Craspedomonadaceae Senn 1900; Craspedophyceae Chadefaud 1960; Craspédomonadophycidées Bourrelly 1968; Craspedomonadophyceae Hibberd 1976; Choanomonadea Krylov et al. 1980; Choanoflagelliida Lee, Hutner & Bovee 1985; Choanoflagellatea Cavalier-Smith 1997 emend. Cavalier-Smith 1998; Choanomonada Adl et al. 2005].
        • Craspedida Cavalier-Smith 1997, emend. Nitsche et al. 2011
        • Acanthoecida Cavalier-Smith 1997, emend. Nitsche et al. 2011
          • Acanthoecidae Norris 1965, emend. sensu Nitsche et al. 2011. Acanthoeca, Enibas,[20] Helgoeca, Polyoeca, Savillea.
          • Stephanoecidae Leadbeater 2011. Acanthocorbis, Amoenoscopa, Apheloecion, Bicosta, Calliacantha, Calotheca, Campanoeca, Campyloacantha, Conion, Cosmoeca, Crinolina, Crucispina, Diaphanoeca, Didymoeca, Kakoeca, Monocosta, Nannoeca, Parvicorbicula, Platypleura, Pleurasiga, Polyfibula, Saepicula, Saroeca, Spinoeca, Spiraloecion, Stephanacantha, Stephanoeca, Syndetophyllum.
      • Metazoa Haeckel 1874, emend. Adl et al. 2005 [Animalia Linnaeus 1758; Eumetazoa Bütschli 1910]. Excluded from protists.

Amoebozoa Lühe 1913, sensu Cavalier-Smith 1998

Cladogram of Amoebozoa based on an 824-gene phylogenetic analysis published in 2022.[21]

Incertae sedis Amoebozoa: Belonocystis, Boveella, Biomyxa, Corallomyxa, Gibbodiscus, Hartmannia, Malamoeba, Malpighamoeba, Microglomus, Oscillosignum, Pseudothecamoeba, Rhabdamoeba, Schoutedamoeba, Stereomyxa, Subulamoeba, Thecochaos, Triaenamoeba, Unda.[3]

Tubulinea Smirnov et al. 2005

Evosea Kang et al. 2017

Discosea Cavalier-Smith et al. 2004 sensu Smirnov et al. 2011

Diaphoretickes Adl et al. 2012

Archaeplastida Adl et al. 2005

  • Glaucophyta Skuja 1954 [Glaucocystaceae West 1904; Glaucocystophyta Kies and Kremer 1986].[5] Incertae sedis Glaucophyta: ?Archaeopsis, ?Chalarodora, ?Glaucocystopsis, ?Peliaina, ?Strobilomonas.
  • Picozoa Seenivasan et al. 2013 [Picobiliphytes Not et al. 2007]. Picomonas.[n]
  • Rhodelphidia Gawryluk et al. 2019. Rhodelphis.[37]
  • Rhodophyceae Thuret 1855, emend. Rabenhorst 1863 [Rhodophyta Wettstein 1901; Rhodoplantae Saunders and Hommersand 2004 emend. Adl et al. 2005]
  • Chloroplastida Adl et al. 2005 [Viridiplantae Cavalier-Smith 1981; Chlorobionta Jeffrey 1982, emend. Bremer 1985, emend. Lewis and McCourt 2004; Chlorobiota Kendrick and Crane 1997]

Stramenopiles Patterson 1989, emend. Adl et al. 2005

Stramenopiles incertae sedis:

  • Environmental clades MAST-21, MAST-25 Massana et al. 2014
  • Platysulcidae Shiratori, Nakayama & Ishida 2015. Platysulcus.
  • Kaonashiidae Weston, Eglit & Simpson 2023. Kaonashia.[38]

Bigyra Cavalier-Smith 1998, emend. 2006

Gyrista Cavalier-Smith 1998

Alveolata Cavalier-Smith 1991

Palustrimonas

Ciliophora

Acavomonas

Colponema

Loeffela

Cladogram of Alveolata, based on a SSU rDNA phylogeny of basal alveolates[53] and a 203-gene phylogeny of apicomplexans and their relatives[54] published in 2023.
  • Colponemida Cavalier-Smith 1993 emend. Adl et al. 2019 (P)
    • Colponemidia Tikhonenkov et al. 2014. Colponema.
    • Acavomonidia Tikhonenkov et al. 2014. Acavomonas.
    • Neocolponema Gigeroff et al. 2023[53]
    • Loeffela Gigeroff et al. 2023[53]
    • Palustrimonas Patterson & Simpson 1996

Myzozoa Cavalier-Smith & Chao 2004

Ciliophora Doflein 1901

  • Postciliodesmatophora Gerassimova & Seravin 1976
  • Intramacronucleata Lynn 1996. Incertae sedis: Protocruziidae Jankowski 1980 [Protocruziidia de Puytorac et al. 1987] (Protocruzia Faria da Cunha & Pinto 1922).
    • SAL Gentekaki et al. 2014. Incertae sedis: 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)[r]
              • 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: 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: 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: 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: 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,[73] 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.[74]
                • 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: 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: 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: 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.

Rhizaria Cavalier-Smith 2002

Rhizaria incertae sedis: Gymnosphaerida Poche 1913, emend. Mikrjukov 2000. Actinocoryne, Cienkowskya, Gymnosphaera, Hedraiophrys [possible synonym of Cienkowskya], Wagnerella.

Cercozoa Cavalier-Smith 1998, emend. Adl et al. 2005; emend. Cavalier-Smith 2018

  • Cercozoa incertae sedis:
    • Aquavolonida Bass & Berney 2018. Aquavolon, Lapot.[75]
    • 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: 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. Chlamydophryse, Diaphoropodon, Lecythium, Trachyrhizium.
      • 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: Phaeobola.[78]
        • Cyphoderiidae de Saedeleer 1934. Campascus*, Corythionella, Cyphoderia [Schaudinnula], Messemvriella*, Pseudocorythion, Ichthyosquama, Knarr, Oleiformis, Psammoderia.[79]
        • 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[s] Deflandre 1928
      • Trivalvulariida Siemensma & Dumack 2020.[80] Leptogromia, Trivalvularis.
  • Metromonadea Cavalier-Smith 2007, emend. Cavalier-Smith 2011. Metromonas, Metopion, Micrometopion, Kiitoksia.
  • Granofilosea Cavalier-Smith & Bass 2009.[81] Incertae sedis granofilosean genera*: ?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 Cavalier-Smith 2002, emend. Bass & Berney in Adl et al. 2019

Retaria Cavalier-Smith 2002

  • Foraminifera d'Orbigny 1826. Incertae sedis groups: ?Lagenida, ?Heterogromia, ?Komokiacea*.
    • Monothalamids Pawlowski et al. 2013 (P) [currently under revision].[89] All genera traditionally in Allogromiida, Astrorhizida, and the Xenophyophorea. Genera unplaced into families: Allogromia, Astrammina, Crithionina, Notodendrodes, Psammophaga, Bathysiphon.
      • Nujappikia Gooday & Holzmann 2022[90]
      • Allogromiidae Rhumbler 1904. Vellaria.[91]
      • Lieberkuehniidae Siemensma et al. 2021.[92] Lieberkuehnia, Claparedellus.
      • Edaphoallogromiidae Siemensma et al. 2021.[92] Edaphoallogromia.
      • Lacogromiidae Siemensma & Holzmann 2021.[92] Lacogromia.
      • Limnogromiidae Holzmann & Siemensma 2021.[92] Limnogromia.
      • Reticulomyxidae Page 1987, emend. Hülsmann 2014. Reticulomyxa, Dracomyxa, Haplomyxa,[93] Wobo.
      • Schizamminidae Nørvang, 1961. Jullienella, Schizammina, Spiculosiphon.[94]
      • Velamentofexidae Völcker & Clauß 2021.[92] Velamentofex.
    • Tubothalamea Pawlowski et al. 2012. Fusulinida Delage & Hérouard 1896 probably belongs to this group.[3]
    • Globothalamea Pawlowski et al. 2012
      • Rotaliida Delage & Hérouard 1896
        • Planorbulinidae Schwager 1877. Planorbulinella, Hyalinea.
        • Discorboidea Ehrenberg 1838
          • Discorbidae Ehrenberg 1838. Discorbis.
          • Rosalinidae Reiss 1963. Rupertina, Discanomalina, "Rosalina", Gavelinopsis, Planorbulina.
        • Rotalioidea Ehrenberg 1839, emend. Pawlowski 2013. Incertae sedis genera: ?Criboelphidium, ?"Elphidium", ?Protelphidium.
          • Elphidiidae Galloway 1933. Elphidium.
          • Ammoniidae Saidova 1981. Ammonia.
          • Elphidiellidae Holzmann & Pawlowski 2017. Elphidiella.
          • Haynesinidae Mikhalevich 2013. Haynesina, Aubignyna.
        • Glabratelloidea Loeblich & Tappan 1964
          • Rotaliellidae Loeblich & Tappan 1964. Rotaliella, Rossyatella.
          • Buliminoididae Seiglie 1970. Buliminoides.
          • Glabratellidae Loeblich & Tappan 1964. Glabratella, Glabratellina, Angulodiscorbis, Planoglabratella.
        • Calcarinoidea Schwager 1876
          • Calcarinidae Schwager 1876. Neorotalia, Baculogypsina, Baculogypsinoides, Schlumbergerella, Pararotalia.
        • Nummulitoidea de Blainville 1827
          • Nummulitidae de Blainville 1827. Nummulites, Operculinella, Cycloclypeus, Heterostegina, Operculina, Planoperculina, Planostegina.
        • Serioidea Holzmann & Pawlowski 2017
          • Uvigerinidae Haeckel 1894. Uvigerina, Rectuvigerina, Trifarina.
          • Bolivinitidae Glaessner 1937. Bolivina, Brizalina, Saidovina.
          • Cassidulinidae d'Orbigny 1839. Globocassidulina, Cassidulinoides, Evolvocassidulina, Islandiella, Ehrenbergina
          • Sphaeroidinidae Cushman 1927. Sphaeroidina.
          • Globobuliminidae Cushman 1927. Globobulimina.
        • Incertae sedis Rotaliida families:
          • ?Nonionidae Schultze 1854. Nonion, Nonionella, Nonionellina, Nonionoides.
          • ?Virgulinellidae Loeblich & Tappan 1984. Virgulinella.
          • ?Buliminidae Jones 1875. Bulimina.
          • ?Epistominellidae Holzmann & Pawlowski 2017. Epistominella.
          • ?Stainforthiidae Reiss 1963. Stainforthia, Gallitellia.
          • ?Cibicididae Cushman 1927. Cibicides, Cibicidoides, Heterolepa.
          • ?Chilostomellidae Brady 1881. Chilostomella.
          • ?Pullenidae Schwager 1877. Pullenia.
          • ?Nuttalidae Saidova 1981. Nuttalides.
          • ?Discorbinellidae Sigal 1952. Discorbinella, Hanzawaia.
          • ?Astrononionidae Cushman & Edwards 1937. Astrononion.
          • ?Oridorsalidae Loeblich & Tappan 1984. Oridorsalis.
          • ?Melonidae Holzmann & Pawlowski 2017. Melonis.
          • ?Cymbaloporidae Cushman 1927. Cymbaloporella.
          • ?Rubratelliidae Holzmann & Pawlowski 2017. Rubratella.
          • ?Murrayinelliidae Holzmann & Pawlowski 2017. Murrayinella.
      • Globigerinida Lankester 1885 emend. Arz, Arenillas & Gilabert 2022[7] (P)
        • Parvularugoglobigerinoidea Arenillas, Arz & Gilabert 2022
          • Parvularuglobigerinidae Arenillas, Arz & Gilabert 2022. Pseudocaucasina, Palaeoglobigerina, Parvularugoglobigerina.
        • Eoglobigerinoidea BouDagher-Fadel 2012
          • Eoglobigerinidae Blow 1979. Alicantina, Eoclavatorella, Eoglobigerina, Parasubbotina, Pseudoglobigerinella, Subbotina, Turbeogloborotalia.
          • Globoquadrinidae Blow 1979. Dentoglobigerina, Globoquadrina.
          • Neoacarininidae BouDagher-Fadel 2012. Neoacarinina.
          • Porticulasphaeridae Banner 1982. Globigerinatheka, Guembelitrioides, Inordinatosphaera, Orbulinoides, Porticulasphaera.
        • Globigerinoidea Carpenter, Parker & Jones 1862
          • Globigerinidae Carpenter, Parker & Jones 1862. Alloglobigerinoides, Ciperoella, Globigerina, Globicuniculus, Globigerinoides, Globigerinoidesella, Globoturborotalita, Trilobigerina [Trilobatus], Zeaglobigerina.
          • Globigerinellidae BouDagher-Fadel 2012. Beella, Bolliella, Globigerinella, Orcadia, Protentella, Quiltyella.
          • Hastigerinidae Bolli, Loeblich & Tappan 1957. Hastigerina, Hastigerinella, Hastigerinopsis.
          • Orbulinidae Schultze 1854. Biorbulina, Candorbulina, Orbulina, Praeorbulina.
          • Sphaeroidinellidae Banner & Blow 1959. Prosphaeroidinella, Sphaeroidinella, Sphaeroidinellopsis.
          • Turborotalitidae Hofker 1976. Berggrenia, Turborotalita.
        • Globorotalioidea Cushman 1927
          • Catapsydracidae Bolli, Loeblich & Tappan 1957. Catapsydrax, Globigerinopsoides, Globigerinoita, Velapertina.
          • Globorotaloididae Banner & Blow 1959. Clavatorella, Globorotaloides, Protentelloides.
          • Globorotaliidae Cushman 1927. Dentigloborotalia, Globoconella, Globorotalia, Hirsutella, Jenkinsella, Menardella, Neogloboquadrina, Paragloborotalia, Truncorotalia.
          • Pulleniatinidae Cushman 1927. Pulleniatina.
        • Globanomalinidoidea Loeblich & Tappan 1984
          • Globanomalinidae Loeblich & Tappan 1984. Carinoturborotalia, Globanomalina, Luterbacheria, Planoglobanomalina, Turborotalia.
        • Hantkeninoidea Cushman 1927
          • Pseudohastigerinidae Canudo 1990. Pseudohastigerina.
          • Hantkeninidae Cushman 1927. Applinella, Aragonella, Clavigerinella, Cribrohantkenina, Hantkenina.
        • Truncorotaloidinoidea BouDagher-Fadel 2012
          • Globigerapsidae Blow 1979. Globigerapsis, Muricoglobigerina.
          • Planorotalitidae BouDagher-Fadel 2012. Astrorotalia, Planorotalites.
          • Truncorotaloididae Loeblich & Tappan 1961. Acarinina, Igorina, Morozovella, Morozovelloides, Pearsonites, Praemurica, Pseudogloboquadrina, Testacarinata, Truncorotaloides.
      • Globotruncanida Arz, Arenillas & Gilabert 2022[7]
        • Abathomphaloidea Arz, Arenillas & Gilabert 2022
          • Abathomphalidae Pessagno 1967. Abathomphalus.
          • Globotruncanellidae Maslakova 1964. Globotruncanella, Spinoglobotruncanella.
        • Favuselloidea Longoria 1974
          • Conoglobigerinidae BouDagher-Fadel, Banner & Whittaker 1997. Conoglobigerina, Tenuigerina, ?Sphaerogerina.
          • Favusellidae Longoria 1974. Ascoliella, Favusella, Koutsoukosia.
          • Globuligerinidae Loeblich & Tappan 1984. Compactogerina, Globuligerina, Haeuslerina, Petaloglobigerina.
        • Hedbergelloidea Longoria & Gamper 1975
          • Ananiidae El-Nakhal 2010. Anania, Badriella, Costellagerina, Hillsella, Liuenella [Liuella], Loeblichella, Muricohedbergella, Paracostellagerina, Pessagnoina, Planohedbergella, Pseudoclavihedbergella, Vanhintella.
          • Hedbergellidae Loeblich & Tappan 1961. Asterohedbergella, Clavihedbergella, Hedbergella, Microhedbergella, Paraticinella, Pseudoguembelitria.
          • Helvetoglobotruncanidae Lamolda 1976. Angulocarinella, Bermudeziana, Bollitruncana, Fingeria, Hedbergellita, Brittonella, Helvetoglobotruncana, Unitruncatus, Whiteinella.
          • Praehedbergellidae Banner & Desai 1988. Blefuscuiana, Gorbachikella, Praehedbergella, Lilliputianella, Lilliputianelloides, Wondersella.
        • Globotruncanoidea Brotzen 1942
          • Globotruncanidae Brotzen 1942. Contusotruncana, Globotruncana, Marginotruncana, Obliquacarinata, Sphaerotruncana, Ventrotruncana.
          • Praeglobotruncanidae Ion 1983. Concavatotruncana, Dicarinella, Falsotruncana, Rotundina, Praeglobotruncana, Verotruncana.
          • Reissidae Korchagin 2001. Elevatotruncana, Globotruncanita, Kassabiana, Radotruncana, Sigalitruncana, Turbotruncana.
        • Planomalinoidea Bolli, Loeblich & Tappan 1957
          • Eohastigerinellidae Loeblich & Tappan 1984. Eohastigerinella, Hastigerinoides.
          • Globigerinelloididae Longoria 1974. Alanlordella, Allotheca, Biglobigerinella, Blowiella, Claviblowiella, Globigerinelloides, Pseudoschackoina.
          • Planomalinidae Bolli, Loeblich & Tappan 1957. Bannerina, Planomalina, Pseudoplanomalina.
          • Schackoinidae Pokorný 1958. Asymetria, Groshenyia, Leupoldina, Neoschackoina, Schackoina.
        • Rotaliporoidea Sigal 1958
          • Rotaliporidae Sigal 1958. Anaticinella, Pseudothalmanninella, Rotalipora, Thalmanninella.
          • Ticinellidae Longoria 1974. Biticinella, Claviticinella, Ticinella.
        • Rugoglobigerinoidea Subbotina 1959
          • Helvetiellidae Longoria & Gamper 1984. Archaeoglobigerina, Bucherina, Dorbignya, Edgarinella, Gandolfia, Gansserina, Globocarinata, Helvetiella, Kuglerina.
          • Rugoglobigerinidae Subbotina 1959. Archaeoglobitruncana, Plummerita, Rugoglobigerina, Rugosocarinata, Rugotruncana, Trinitella.
      • Heterohelicida Fursenko 1958, emend. Arz, Arenillas & Gilabert 2022[7]
        • Heterohelicoidea Cushman 1927
          • Gublerinidae Aliyulla 1977. Gublerina, Lipsonia, Praegublerina.
          • 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.
          • Pseudotextulariidae Maamouri & Salaj 1978. Planoglobulina, Pseudotextularia, Racemiguembelina.
          • Pseudoguembelinidae Aliyulla 1977. Eicheriella, Leptobimodalia, Pseudoguembelina.
          • Spiroplectidae Cushman 1911. Hendersonites [Hendersonia], Neohendersonites, Paraspiroplecta, Spiroplecta.
          • Ventilabrellidae Maamouri & Salaj 1978. Planulitella, Proliferania, Sigalia, Ventilabrella.
      • Guembelitriida Arenillas, Arz & Gilabert 2022[7]
        • Cassigerinelloidea Bolli, Loeblich & Tappan 1957
        • Guembelitrioidea Montanaro-Gallitelli 1957
          • Guembelitriidae Montanaro-Gallitelli 1957. Cassigerinelloita, Guembelitria, Chiloguembelitria, Jenkinsina, ?Archaeoguembelitria, ?Gallitellia, ?Guembelitriella.
        • Chiloguembelinoidea Reiss 1963
          • Chiloguembelinidae Reiss 1963. Chiloguembelina, Woodringina, ?Streptochilus.
        • Globoconusoidea BouDagher-Fadel 2012
          • Globoconusidae BouDagher-Fadel 2012. Globoconusa, Trochoguembelitria,
      • Order ? (unnamed)[7]
        • Globigerinitoidea BouDagher-Fadel 2012, emend. Arenillas, Arz & Gilabert 2022[7]
          • Candeinidae Cushman 1927. Candeina.
          • Globigerinatellidae BouDagher-Fadel 2012. Globigerinatella.
          • Globigerinitidae Bermúdez 1961. Globigerinita, Mutabella, Polyperibola, Tinophodella.
          • Tenuitellidae BouDagher-Fadel 2012. Praetenuitella, Tenuitella, Tenuitellinata, Tenuitellita.
          • Family ? (unnamed). Dipsidripella, ?Antarcticella.
      • Robertinida Loeblich & Tappan 1984. Hoeglundina, Robertina, Robertinoides.
      • Textulariida Delage & Hérouard 1896 (P). Cyclammina, Eggerella, Reophax, Textularia, Trochammina.
      • Carterinida Loeblich & Tappan 1981. Carterina.
  • 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 1838, emend. Haeckel 1887
      • Spumellaria Ehrenberg 1875, Haeckel 1887, emend. Riedel 1967. Actinomma, Didymocyrtis, Euchitonia, Hexacontium, Hexalonche, Hexastylus, Octodendron, Plegmosphaera, Saturnalis, Spongaster, Spongosphaera.
      • 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[95]
        • 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[t]
        • Entactiniidae Riedel 1967. Afanasievella, Apophysiactinia, Borisella, Callela, Cancellosphaera, Costaentactinia, Cyclocarpus nomen dubium, Duodecimentactinia, Langsonsphaera,[96] 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.[97]
        • Tetrentactiniidae Kozur & Mostler 1979. Multisphaera.
        • Triassothamnidae Takahashi, Maekawa & Dumitrica 2022. Triassothamnus.[96]

Haptista Cavalier-Smith 2003

  • 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 Shishkin & Zlatogursky 2018. Spiculophrys.
    • Pterocystida Cavalier-Smith and von der Heyden 2007, emend. Shishkin and Zlatogursky 2018
      • Raphidista Shishkin & Zlatogursky 2018
        • Choanocystidae Cavalier-Smith & von der Heyden 2007. Choanocystis.
        • Raphidiophryidae Febvre-Chevalier & Febvre 1984, emend. Shishkin & Zlatogursky 2018. Raphidiophrys.
      • Pterista Shishkin & Zlatogursky 2018
        • Oxnerellidae Cavalier-Smith & Chao 2012. Oxnerella.
        • Pterocystidae Cavalier-Smith & von der Heyden 2007. Pterocystis, Raineriophrys, Chlamydaster, Pseudoraphidiophrys, Pseudoraphidocystis, Triangulopteris.[99]
        • Heterophryidae Poche 1913, emend. Cavalier-Smith & von der Heyden 2007. Heterophrys, Sphaerastrum.
    • Panacanthocystida Shishkin & Zlatogursky 2018
      • Yogsothothidae Shishkin & Zlatogursky 2018 [Chthonida Shishkin & Zlatogursky 2018; Yogsothothina Shishkin & Zlatogursky 2018]. Yogsothoth.
      • Acanthocystida Cavalier-Smith & von der Heyden 2007 emend. Shishkin & Zlatogursky 2018
        • Marophryidae Cavalier-Smith & von der Heyden 2007 emend. Shishkin & Zlatogursky 2018 [Marophryina Cavalier-Smith in Yabuki et al. 2012]. Marophrys.
        • Chalarothoracina Hertwig & Lesser 1874 sensu Cavalier-Smith in Yabuki et al. 2012 emend. Shishkin & Zlatogursky 2018
          • Acanthocystidae Claus 1874 emend. Shishkin & Zlatogursky 2018. Acanthocystis.
          • Raphidocystidae Zlatogursky 2018. Raphidocystis.

Cryptista Cavalier-Smith 1989, 2018, emend. Adl et al. 2019

Incertae sedis Eukarya

Hemimastigophora Foissner et al. 1988

Excavates (P)

Malawimonadida Cavalier-Smith 2003

Metamonada Grassé 1952, emend. Cavalier-Smith 1987

  • ?Barthelona
  • Anaeramoebae[103] [Anaeramoebidae Táborský, Pánek & Čepička 2017]. Anaeramoeba.[104]
  • Fornicata Simpson 2003
  • Parabasalia Honigberg 1973
    • ?Tricercomitus
    • Trichomonadida Kirby 1947. Cochlosoma, Dientamoeba, Lacusteria, Pentatrichomonas, Pentatrichomonoides, Pseudotrichomonas, Pseudotrypanosoma, Tetratrichomonas, Trichomonas, Trichomonoides, Trichomitopsis.
    • Honigbergiellida Čepička et al. 2010 (P?). Ditrichomonas, Cthulhu, Cthylla, Hexamastix, Honigbergiella, Monotrichomonas.
    • Hypotrichomonadida Čepička et al. 2010. Hypotrichomonas, Trichomitus.
    • Tritrichomonadida Čepička et al. 2010 (P?). Dientamoeba, Histomonas, Monocercomonas, Parahistomonas, Simplicimonas, Tritrichomonas.
    • Cristamonadida Brugerolle & Patterson 2001. Caduceia, Calonympha, Coronympha, Deltotrichonympha, Devescovina, Foaina, Gigantomonas, Joenia, Joenina, Joenoides, Joenopsis*, Kofoidia, Koruga, Macrotrichomonas, Macrotrichomonoides, Metadevescovina, Mixotricha, Pachyjoenia*, Projoenia*, Pseudodevescovina, Rhizonympha*, Snyderella, Stephanonympha.
    • Spirotrichonymphida Grassé 1952. Holomastigotes*, Holomastigotoides, Microjoenia*, Micromastigotes*, Rostronympha*, Spiromastigotes*, Spironympha*, Spirotrichonympha*, Spirotrichonymphella, Uteronympha*.
    • Lophomonadida Light 1927. Lophomonas.
    • Trichonymphida Poche 1913. Barbulanympha, Eucomonympha, Heliconympha, Hoplonympha, Leptospironympha, Macrospironympha*, Pseudotrichonympha, Rhynchonympha*, Spirotrichosoma*, Staurojoenina, Teranympha, Trichonympha, Urinympha.
  • Preaxostyla Simpson 2003

Discoba Simpson in Hampl et al. 2009

CRuMs Brown et al. 2018

  • Collodictyonidae Brugerolle et al. 2002, emend. Adl et al. 2019 [Diphylleidae Cavalier-Smith 1993; Diphylleida Cavalier-Smith 1993; Diphyllatea Cavalier-Smith 2003, Sulcomonadidae Cavalier-Smith 2013]. Collodictyon, Diphylleia [Aulacomonas], Sulcomonas.
  • Rigifilida Cavalier-Smith in Yabuki et al. 2012 [Hilomonadea Cavalier-Smith 2008 emend. 2012]
  • Mantamonadidae Cavalier-Smith in Glücksman et al. 2011 [Glissodiscea Cavalier-Smith 2013 emend. 2021; Mantamonadida Cavalier-Smith in Glücksman et al. 2011]. Mantamonas.

Ancyromonadida Cavalier-Smith 1998

  • Ancyromonadidae Cavalier-Smith 2008. Ancyromonas, Nutomonas.
  • Planomonadidae Cavalier-Smith 1993. Fabomonas, Planomonas.

Genera of uncertain affiliation

Acinetactis, Actinastrum, Actinocoma, Actinolophus, Adinomonas, Aletium, Amphimonas, Amylophagus, Aphelidiopsis, Asterocaelum, Asthmatos, Aurospora, Barbetia, Berkeleyaesol,[118] Belaria, Belonocystis, Bertarellia, Bertramia, Bodopsis, Boekelovia, Branchipocola, Camptoptyche, Chalarodora, Cibdelia, Cichkovia, Cinetidomyxa, Cingula, Cladomonas, Clathrella, Codonoeca, Coelosporidium[u], 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,[119] Gymnococcus, Gymnophrydium, Haematotractidium, Hartmannina, Heliobodo, Heliomonas, Hermisenella, Heterogromia, Hillea, Hyalodaktylethra, Immanoplasma, Isoselmis, Janickina, Kamera, Lagenidiopsis, Liegeosia, Luffisphaera[v], Lymphocytozoon, Lymphosporidium, Macappella, Magosphaera, Malpighiella, Martineziella, Megamoebomyxa, Meringosphaera, Microcometes, Monochrysis, Monodus, Mononema, Myrmicisporidium, Naupliicola, Nephrodinium, Neurosporidium, Orbulinella, Ovicola, Palisporomonas, Pansporella, Paradinemula, Parakaryon[w], 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, Tetragonidium, Thaulirens, Topsentella, Toshiba, Trichonema, Urbanella.[3]

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. ^ a b This taxon belongs to the paraphyletic phylum Apusozoa Cavalier-Smith 1997 emend. 2013.[15]
  4. ^ a b c There are two competing phylogenetic hypotheses: Teretosporea (Corallochytrium + Ichthyosporea) and Pluriformea (Corallochytrium + Syssomonas).[3] This article follows the latter hypothesis, because it is strongly supported by the most recent phylogenetic studies.[11]
  5. ^ The term "Choanozoa" has been used since 1991 by Cavalier-Smith as a paraphyletic assemblage of opisthokont protists, and the terms "Apoikozoa" and "choanimal" were proposed as names for the clade Metazoa+Choanoflagellata. However, these terms have not been formally described or adopted, and were rejected in favour of a renamed Choanozoa to fit the clade Metazoa+Choanoflagellata.[3]
  6. ^ 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.[3]
  7. ^ The latest revision by Adl et al. (2019) ignores the grouping of some variosean genera into higher rank clades that was made in older studies because of the weakly supported SSU phylogenetic analyses, except for Fractovitellida, which is fully supported by phylogenomic analyses from 2017.[3]
  8. ^ a b c d e f g This order was 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.[22][3]
  9. ^ This genus groups with Dermamoebidae in a phylogenetic anaysis that uses a limited number of taxa.[30] In the recent taxonomy it is listed as a separate clade until its position is better resolved.[3]
  10. ^ This genus could be a synonym of Vannella.[3]
  11. ^ Both genera group together in some phylogenetic analyses, but usually appear separated. They are considered a potential branch of Discosea, unless proven otherwise in the future with better taxon sampling.[3]
  12. ^ Latest analyses show that Microheliella is the sister group to Cryptista.[12]
  13. ^ Telonemia is the sister group to the SAR clade.[9]
  14. ^ The picobiliphytes are phylogenetically closer to Rhodophyta and Rhodelphis than to other groups, and are therefore considered part of Archaeplastida.[36]
  15. ^ Cavalier-Smith and Scoble (2013) show a sister relationship between raphidophytes and Actinophryida.[47]
  16. ^ Apart from morphological descriptions, no molecular analysis has solidly affiliated Rastrimonas with the Perkinsozoa.[58]
  17. ^ Highly divergent 18S rRNA.[3]
  18. ^ This taxon is artificial; many of the families listed in it are not monophyletic and have little support from phylogenetic analyses.[3]
  19. ^ There is only one molecular sequence from this genus, which causes long branches in gene phylogenies, and the node is unresolved.[3]
  20. ^ 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.[95]
  21. ^ Probably a synonym of the zygomycete fungus Nephridiophaga.[3]
  22. ^ May be the same genus as Belonocystis.[3]
  23. ^ It is not yet known whether this organism is an eukaryote or a prokaryote.[120]

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