Classification of eukaryotes
A modern non-hierarchical interpretation of the phylogeny of eukaryotes, including protists
A protist () 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 taxon.
† = Extinct taxon.
? = Taxon of uncertain taxonomic position .
* = Taxon lacking molecular data.
[ ] = Synonyms and alternative names.
→ = Monotypic taxon.
Current classification [ edit ]
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]
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]
Nucletmycea Brown et al. 2009 [Holomycota Liu et al. 2009 ]
Rotosphaerida Rainer 1968 [Cristidiscoidida Page 1987, Cavalier-Smith 1993, 1998 ; Nucleariidae Patterson 1983, 1999 ]. Fonticula , Nuclearia , Parvularia , Pompholyxophrys , Lithocolla , ?Vampyrellidium *, ?Elaeorhanis *, ?Pinaciophora *, ?Rabdiophrys *, ?Rabdiaster *, ?Thomseniophora *.[ 17]
Fungi R.T. Moore 1980 . Excluded from protists.
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 ]
Dermocystida Cavalier-Smith 1998 [Rhinosporidaceae Mendoza et al. 2001 ]. Amphibiocystidium , Amphibiothecum , Chromosphaera , Dermocystidium , Rhinosporidium , Sphaerothecum .
Ichthyophonida Cavalier-Smith 1998 [Ichthyophonae Mendoza et al. 2001 ; Amoebidiidae Reeves 2003 ]. Abeoforma , Amoebidium , Anurofeca , Astreptonema , Caullerya , Creolimax , Eccrinidus , Enterobryus , Enteropogon , Ichthyophonus , Palavascia , Pseudoperkinsus , Psorospermium , Sphaeroforma .
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
Salpingoecidae Kent 1880–1882, emend. sensu Nitsche et al. 2011 . Astrosiga , Aulomonas , Barroeca ,[ 20] Choanoeca , Cladospongia , Codonocladium , Codonosigopsis , Codosiga [Codonosiga ], Desmarella [Codonodesmus ; Kentrosiga ], Dicraspedella , Diploeca , Diplosiga , Diplosigopsis , Hartaetosiga , Kentia , Lagenoeca , Microstomoeca , Monosiga , Mylnosiga , Pachysoeca , Proterospongia , Salpingoeca , Salpingorhiza , Sphaeroeca , Stagondoeca , Stelexomonas , Stylochromonas .
Acanthoecida Cavalier-Smith 1997, emend. Nitsche et al. 2011
Acanthoecidae Norris 1965, emend. sensu Nitsche et al. 2011 . Acanthoeca , Enibas ,[ 21] 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.
Cladogram of Amoebozoa based on an 824-gene phylogenetic analysis published in 2022.[ 22]
Incertae sedis Amoebozoa: Belonocystis , Boveella , Biomyxa , Corallomyxa , Gibbodiscus , Hartmannia , Malamoeba , Malpighamoeba , Microglomus , Oscillosignum , Pseudothecamoeba , Rhabdamoeba , Schoutedamoeba , Stereomyxa , Subulamoeba , Thecochaos , Triaenamoeba , Unda .[ 3]
Corycida Kang et al. 2017 ["Corycidia"][ 23]
Echinamoebia Cavalier-Smith et al. 2016 → Echinamoebida Cavalier-Smith et al. 2004 em. Smirnov et al. 2011 .
Elardia Kang et al. 2017 [ 23]
Leptomyxia Cavalier-Smith et al. 2016 → Leptomyxida Pussard & Pons 1976 emend. Page 1987
Eulobosia Cavalier-Smith et al. 2016 :
Arcellinida Kent 1880 . Incertae sedis genera: Acipyxis , Apolimia , Argynnia , Armipyxis , Conicocassis , Cornuapyxis , Ellipsopyxella , Ellipsopyxis , Erugomicula , Frenopyxis , Geamphorella , Geoplagiopyxis , Geopyxella , Hoogenraadia , Jungia , Lagenodifflugia , Lamptopyxis , Lamptoquadrula , Leptochlamys , Maghrebia , Meisterfeldia , Microquadrula , Nabranella , Oopyxis , Paracentropyxis , Paraquadrula , Pentagonia , Physochila , Planhoogenraadia , Pomoriella , Pontigulasia , Prantlitina ,[ 26] Proplagiopyxis , Protoplagiopyxis , Protocucurbitella , Pseudawerintzewia , Pseudonebela , Schoenbornia , Sexangularia , Suiadifflugia , Trigonopyxis , Wailesella .[ 6]
Phryganellina Bovee 1985
Organoconcha Lahr et al. 2019
Glutinoconcha Lahr et al. 2019
Volnustoma Lahr et al. 2019
Hyalospheniformes Lahr et al. 2019
Hyalospheniidae Schultze 1877 emend. Kosakyan et al. 2012 . Alabasta , Alocodera , Apodera , Certesella , Cornutheca , Gibbocarina , Hyalosphenia , Longinebela , Mrabella , Nebela , Padaungiella , Planocarina , Porosia , Quadrulella .
Excentrosoma Lahr et al. 2019
Cylindrothecina González-Miguéns, Todorov, Blandenier, Porfirio-Sousa, Ribeiro, Ramos, Lahr & Lara 2022
Longithecina Lahr et al. 2019
Sphaerothecina Kosakyan et al. 2016
Euamoebida Lepşi 1960 sensu Smirnov et al. 2011 [ 28]
Amoebina Cavalier-Smith 2016
Amoebidae Ehrenberg 1838 Page 1987 . Amoeba , Chaos , Polychaos , Parachaos , Trichamoeba , Deuteramoeba , Hydramoeba
Hartmannellidae Volkonsky 1931 . Cashia , Copromyxa , Copromyxella , Glaeseria , Hartmannella , Ptolemeba , Saccamoeba
Nolandina Cavalier-Smith 2016 [ 28]
Variosea [ g] Cavalier-Smith 2004
Angulamoeba Berney, Bass & Geisen 2015
Arboramoeba Berney, Bass & Geisen 2015
Darbyshirella Berney, Bass & Geisen 2015
Dictyamoeba Berney, Bass & Geisen 2015
Ischnamoeba Berney, Bass & Geisen 2015
Heliamoeba Berney, Bass & Geisen 2015
Filamoeba Page 1967
Phalansterium Cienkowski 1870 .
Flamellidae Lahr & Katz in Lahr et al. 2011 .[ 29] Flamella , Telaepolella .
Protosteliida Olive & Stoianovitch 1966, sensu Shadwick & Spiegel in Adl et al. 2012 . Protostelium [incl. Planoprotostelium ].
Fractovitellida Lahr et al. 2011, sensu Kang et al. 2017
Cavosteliida Shadwick & Spiegel in Adl et al. 2012 . Cavostelium , Schizoplasmodiopsis , Tychosporium .
Holomastigida Lauterborn 1895 . Artodiscus , Multicilia .
Eumycetozoa Zopf 1884 sensu Kang et al. 2017 [Mycetozoa de Bary 1859 ex Rostafinski 1873 ; Macromycetozoa Fiore-Donno 2010 ]
Dictyostelia Lister 1909 sensu Sheikh et al. 2018 [Dictyosteliomycetes Doweld 2001 ]. Acytostelium , Cavenderia , Coremiostelium , Dictyostelium , Heterostelium , Polysphondylium , Raperostelium , Rostrostelium , Speleostelium , Tieghemostelium , Synstelium , ?Coenonia .
Protosporangiida Shadwick & Spiegel in Adl et al. 2012 [Ceratiomyxea Leontyev, Stephenson, Novozhilov & Shchepin 2019 ; Ceratiomyxomycetes D. Hawksw., B. Sutton & Ainsw. 2019 ][ 4]
Myxogastria Macbride 1899 [Myxogastrea Cavalier-Smith 1993 ; Myxomycetes Link 1833, sensu Haeckel 1866 ]
Lucisporomycetidae [Lucisporinia Leontyev, Stephenson, Novozhilov & Shchepin 2019 ;[ 4] Lucisporidia Cavalier-Smith 2013 ]. Incertae sedis genera: Arcyriatella , Calonema , Minakatella , Trichioides .
Columellomycetidae [Collumellinia Leontyev, Stephenson, Novozhilov & Shchepin 2019 ;[ 4] Columellidia Cavalier-Smith 2015 ]. Incertae sedis genera: Diachea , Echinosteliopsis , Leptoderma , Paradiachea , Protophysarum , Trabrooksia , Willkommlangea .
Echinosteliidia Leontyev, Stephenson, Novozhilov & Shchepin 2019 [ 4]
Stemonitidia Leontyev, Stephenson, Novozhilov & Shchepin 2019 [ 4]
Clastodermatales Leontyev, Stephenson, Novozhilov & Shchepin 2019 [ 4]
Meridermatales Leontyev, Stephenson, Novozhilov & Shchepin 2019 [ 4]
Stemonitidales T. Macbr. 1922 em. Leontyev, Stephenson, Novozhilov & Shchepin 2019
Stemonitidaceae Fr. 1829 em. Leontyev, Stephenson, Novozhilov & Shchepin 2019 . Macbrideola , Stemonitis , Symphytocarpus .
Amaurochaetaceae Rostaf. ex Cooke 1877 em. Leontyev, Stephenson, Novozhilov & Shchepin 2019 . Comatricha , Stemonaria , Stemonitopsis , Amaurochaete , Brefeldia , Enerthenema , Paradiacheopsis .
Physarales T. Macbr. 1922
Tasmaniomyxa umbilicata Lloyd et al. 2023 [ 30]
Lamprodermataceae T. Macbr. 1899 em. Leontyev, Stephenson, Novozhilov & Shchepin 2019 . Lamproderma [incl. Dichaeopsis , Colloderma , Elaeomyxa ], ?Collaria .
Didymiaceae Rostaf. ex Cooke 1877 – Diderma , Didymium , Lepidoderma , Mucilago .
Physaraceae Chevall. 1826 . Craterium , Leocarpus , Fuligo , Physarum , Physarella , Physarina , Kelleromyxa , Badhamia .
Cutosea Cavalier-Smith 2016 [Squamocutida Cavalier-Smith 2016 ]
Archamoebea Cavalier-Smith 1983, sensu Cavalier-Smith 2004
Discosea Cavalier-Smith et al. 2004 sensu Smirnov et al. 2011 [ edit ]
Flabellinia Smirnov et al. 2005
Dermamoebida Cavalier-Smith 2004
Thecamoebida [ h] Schaeffer 1926, sensu Smirnov & Cavalier-Smith 2011 . Sappinia , Stenamoeba , Stratorugosa ,[ 34] Thecamoeba , Thecochaos .[ 35]
Dactylopodida [ h] Smirnov et al. 2005 sensu Kang et al. 2017 . Cunea , ?Janickina ,* Korotnevella , Neoparamoeba , Paramoeba , Pseudoparamoeba , Vexillifera .
Vannellida Smirnov et al. 2005 [Vannellidae Bovee 1979 ]. Clydonella , Lingulamoeba , Paravannella , Pessonella *,[ j] Ripella , Vannella .
Stygamoebida [ k] Smirnov & Cavalier-Smith in Smirnov et al. 2011 (P). Stygamoeba , Vermistella .
Centramoebia Cavalier-Smith et al. 2016 [ 28]
Acanthopodida [ h] Page 1976 [Centramoebida Rogerson & Patterson 2002, sensu Tice et al. 2016 [ 36] ]. Acanthamoeba , Balamuthia , Luapeleamoeba , Protacanthamoeba , Dracoamoeba , Vacuolamoeba .
Himatismenida [ h] Page 1987 . Cochliopodium , Ovalopodium , Parvamoeba .
Pellitida [ h] Smirnov & Cavalier-Smith 2011, sensu Kang et al. 2017 . Endostelium , Gocevia , Paragocevia *, Pellita .
Glaucophyta Skuja 1954 [Glaucocystaceae West 1904 ; Glaucocystophyta Kies and Kremer 1986 ; Glaucoplantae Marin & Melkonian in Li et al. 2020 ]. Incertae sedis taxa: Archaeopsis , Chalarodora , Glaucocystopsis , Peliaina , Strobilomonas .[ 5]
Picozoa Seenivasan et al. 2013 [Picobiliphytes Not et al. 2007 ]: Picomonas .[ n]
Rhodelphidia Tikhonenkov, Gawryluk, Mylnikov & Keeling 2019 : Rhodelphis Tikhonenkov, Gawryluk, Mylnikov & Keeling 2019 .[ 39]
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 ]
Prasinodermophyta Marin & Melkonian in Li et al. 2020 [ 40]
Prasinodermophyceae Marin & Melkonian in Li et al. 2020 → Prasinodermales Marin & Melkonian in Li et al. 2020 → Prasinodermaceae Marin & Melkonian in Li et al. 2020 : Prasinoderma Hasegawa & Chihara in Hasegawa et al. 1996 .
Palmophyllophyceae Leliaert et al. 2016 emend. Marin & Melkonian in Li et al. 2020
Palmophyllales Zechman, Verbruggen, Leliaert, Ashwort, M.A.Buchheim, Fawley, H.Spalding, Pueschel, J.A.Buschheim, Verghese & Hanisak 2010 → Palmophyllaceae Zechman, Verbruggen, Leliaert, Ashwort, M.A.Buchheim, Fawley, H.Spalding, Pueschel, J.A.Buschheim, Verghese & Hanisak 2010 : Palmophyllum Kützing 1849 , Verdigellas Ballantine & Norris 1994 , Palmoclathrus ? Womersley 1971 .[ 41]
Prasinococcales Guillou et al. ex Leliaert et al. 2016 emend. Marin & Melkonian in Li et al. 2020 → Prasinococcaceae Leliaert et al. 2016 : Prasinococcus Miyashita & Chihara in Miyashita et al. 1993 .
Chlorophyta Pascher 1914 emend. Lewis & McCourt 2004
Streptophyta Bremer & Wanntorp 1981 [Charophyta Migula 1897 emend. Karol et al. 2009 ]
Chlorokybophyceae Irisarri et al. 2021 → Chlorokybales Irisarri et al. 2021 → Chlorokybaceae Irisarri et al. 2021 : Chlorokybus Geitler 1942 .[ 42]
Mesostigmatophyceae Marin & Melkonian 1999 emend. Lewis & McCourt 2004 [Mesostigmata Turmel et al. 2002 ] → Mesostigmales Cavalier-Smith 1998 → Mesostigmataceae Fott 1947 : Mesostigma Lauterborn 1894 .
Klebsormidiophyceae C. Jeffrey ex Guiry 2023 [ 43]
Phragmoplastophyta Lecointre & Guyander 2006
Charophyceae Smith 1938 emend. Karol et al. 2009
Coleochaetophyceae Jeffrey 1982
Anydrophyta Rensing 2020 [ 44]
Zygnematophyceae van den Hoek et al. 1995 emend. Hall et al. 2009 [ 45]
Serritaeniales S.Hess & J.de Vries 2022 → Serritaeniaceae S.Hess & J.de Vries 2022 : Serritaenia A.Busch & S.Hess 2021
Zygnematales Bessey 1907 emend. Hess & de Vries 2022 → Zygnemataceae Kützing 1843 : Cylindrocystis Meneghini 1848 , Mesotaenium ? Nägeli 1849 , Mougeotia (Roth) C.Agardh 1824 , Zygnema C.Agardh 1817 nom. cons. , Zygnemopsis (Skuja) Transeau 1934 .
Desmidiales Bessey 1907 emend. S.Hess & J.de Vries 2022 → Desmidiaceae Ralfs 1848 : Bambusina Kützing ex Kützing 1849 nom. cons. , Closterium Nitzsch ex Ralfs 1848 , Cosmarium Corda ex Ralfs 1848 , Cosmocladium Brébisson 1856 , Desmidium C.Agardh ex Ralfs 1848 , Euastrum Ehrenberg ex Ralfs 1848 , Micrasterias C.Agardh ex Ralfs 1848 , Netrium (Nägeli) Itzigsohn & Rothe 1856 , Nucleotaenium Gontcharov & Melkonian 2010 , Onychonema G.C.Wallich 1860 , Penium Brébisson ex Ralfs 1848 , Phymatodocis Nordstedt 1878 , Planotaenium (Ohtani) Petlovany & Palamar-Mordvintseva 2009 , Pleurotaenium Nägeli 1849 , Staurastrum Meyen ex Ralfs 1848 , Staurodesmus Teiling 1948 , Xanthidium Ehrenberg ex Ralfs 1848 , etc.
Spirogyrales Clements nom. cons. emend. Hess & de Vries 2022 : Spirogyra Link 1820 nom. cons.
Embryophyta Engler 1886 emend. Lewis & McCourt 2004 [Plantae Haeckel 1866 ] (excluded from protists)
Stramenopiles incertae sedis :
Opalozoa Cavalier Smith 1991, emend. 2006 . Incertae sedis : Cantina , Rictus , environmental clades MAST-16, MAST-22 and MAST-24 Massana et al. 2014 .
Placidozoa Cavalier-Smith 2013
Bicosoecida Grassé 1926, emend. Karpov 1998 . Adriamonas , Anoeca , Bicosoeca , Caecitellus , Cafeteria , Cafileria ,[ 48] Cyathobodo , Filos , Halocafeteria , Nanum , Paramonas , Pseudobodo , Pseudodendromonas , Otto , Regin .[ 49]
Sagenista Cavalier-Smith 1995 . Incertae sedis : environmental clades MAST-4, MAST-7, MAST-8, MAST-9, MAST-10, MAST-11 and MAST-20 Massana et al. 2014 .
Labyrinthulomycetes Dick 2001
Amphitremida Poche 1913, emend. Gomaa et al. 2003 . Amphitrema , Archerella , Diplophrys , Paramphitrema .
Amphifilida Cavalier-Smith 2012 . Amphifila , Fibrophrys , Sorodiplophrys .
"Oblongichytrida " Bennett et al. 2017 . Oblongichytrium .
Labyrinthulida Doffein 1901 . Aplanochytrium , Labyrinthula , Stellarchytrium .
Thraustochytrida Sparrow 1943 . Althornia , Aurantiochytrium , Botryochytrium , Japanochytrium , Monorhizochytrium , Parietichytrium , Phycophthorum ,[ 50] Schizochytrium , Sicyoidochytrium , Thraustochytrium , Ulkenia .
Pseudophyllomitidae Shiratori et al. 2016 [clade MAST-6].[ 51] Pseudophyllomitus .
Gyrista incertae sedis : environmental clades MAST-1, MAST-2 and MAST-23 Massana et al. 2014 .
Pseudofungi Cavalier-Smith 1986 [Heterokontimycotina M.W. Dick 1976 ]
Bigyromonadea Cavalier-Smith 1998 [ 52]
Developea Karpov & Aleoshin 2016 . Cubaremonas , Develocanicus , Develocauda , Developayella , Develorapax , Mediocremonas .
Pirsoniales Cavalier-Smith 1998 emend. Prokina et al. 2024 .[ 53] Bordeauxia , Bullionia , Feodosia , Koktebelia , Noirmoutieria , Pirsonia .
Hyphochytriales Sparrow 1960 . Anisolpidium , Canteriomyces .
Oomycetes Winter 1897, emend. Dick 1976 [Peronosporomycetes Dick 2001 ]. Incertae sedis genera: Atkinsiella , Ciliomyces , Crypticola , Ectrogella , Eurychasma , Halodaphnea , Haliphthoros , Haptoglossa , Lagena , Lagenisma , Olpidiopsis , Pontisma , Pythiella , Rozellopsis , Sirolpidium .
Saprolegnialean lineage Lara in Adl et al. 2019 . Achlya , Aphanomyces , Aplanopsis , Apodachlya , Aquastella , Geolegnia , Leptomitus , Newbya , Pythiopsis , Protoachlya , Salisapilia , Saprolegnia , Thraustotheca .
Peronosporalean lineage Lara in Adl et al. 2019 . Albugo , Bremia , Chlamydomyzium , Halophytopthora , Hyaloperonospora , Lagenidium , Myzocytiopsis , Peronospora , Plasmopara , Pythium , Pseudoperonospora , Phytophthora , Phytopythium , Pustula .
Ochrophyta Cavalier-Smith 1986, emend. Cavalier-Smith & Chao 1996 [Heterokontophyta Guiry, R.A.Andersen & Moestrup 2023 ]
Chrysista Cavalier-Smith 1986 . Incertae sedis : environmental lineages MOCH-3 and MOCH-5 Massana et al. 2014 , Chrysowaernella , Aurearena .
SII clade
Olisthodiscophyceae Barcytė, Eikrem & M. Eliáš 2021 .[ 54] Olisthodiscus .
Pinguiophyceae Kawachi et al. 2003 . Glossomastix , Phaeomonas , Pinguiochrysis , Pinguiococcus , Polypodochrysis .
Limnistia Cavalier-Smith 1996 emend. 2006
Chrysophyceae Pascher 1914 . Incertae sedis : Chryososaccus , Chrysosphera , Cyclonexis , Lygynion , Phaeoplaca .
Eustigmatophyceae Hibberd & Leedale 1971 [ 55]
Eustigmatales Hibberd 1981
Paraeustigmatos Fawley, Nemcová, & Fawley 2019
Eustigmataceae Hibberd 1981 . ?Ellipsoidion , Chlorobotrys , Eustigmatos , Pseudocharaciopsis , Vischeria .
Monodopsidaceae Hibberd 1981 . Microchloropsis , Monodopsis , Nannochloropsis , Pseudotetraedriella .
Neomonodaceae Amaral et al. 2020 . ?Botryochloropsis , Characiopsella , Munda , Neomonodus , Pseudellipsoidion .
Goniochloridales Fawley, Elias & Fawley 2013 . Goniochloris , Pseudostaurastrum , Tetraedriella , Trachydiscus , Vacuoliviride .
Picophagea Cavalier-Smith 2006 emend. 2017 [ 56] [Synchromophyceae S.Horn & C.Wilhelm 2007 ]. Chlamydomyxa , Chrysopodocystis , Guanochroma , Leukarachnion , Picophagus , Synchroma .
SI clade
Raphidomonadea Chadefaud 1950 ex Silva 1980
PX clade
Chrysomerophyceae Cavalier-Smith 1995 . Antarctosaccion , Chrysomeris , Chrysowaernella , Giraudyopsis , Rhamnochrysis , Tetrasporopsis .
Chrysoparadoxophyceae Wetherbee et al. 2019 .[ 58] Chrysoparadoxa .
Phaeophyceae Hansgirg 1886 [Kjellman 1891; Pfitzer 1894]
Ascoseirales Petrov 1964 . Ascoseira .
Desmarestiales Setchell & Gardner 1925 . Arthrocladia , Desmarestia (P), Himantothallus , Phaeurus .
Dictyotales Bory de Saint-Vincent 1828 . Dictyota , Dilophus , Lobophora , Padina , Stypopodium , Taonia , Zonaria .
Discosporangiales Kawai et al. 2007 . Choristocarpus , Discosporangium .
Ectocarpales Bessey 1907, emend. Silva & Reviers 2000 . Adenocystis , Acinetospora , Asterocladon , Asteronema , Chordaria , Ectocarpus , Scytosiphon .
Fucales Bory de Saint-Vincent 1927 . Ascophyllum , Bifurcaria , Cystoseira , Druvillaea , Fucus , Hormosira , Sargassum , Turbinaria .
Ishigeales Cho et al. 2004 [Ishigeacea Okamura 1935 ]. Ishige .
Laminariales Migula 1908 . Akkesiophycus , Alaria , Chorda , Costaria , Laminaria , Lessonia , Pseudochoda .
Nemodermatales Parente et al. 2008 . Nemoderma .
Onslowiales Draisma & Prud'homme van Reine 2008 . Onslowia .
Ralfsiales Nakamura ex Lim & Kawai 2007 . Lithoderma , Neoralfsia , Pseudolithoderma , Ralfsia .
Scytothamnales Peters & Clayton 1998 . Scytothamnus , Splachnidium , Stereocladon .
Sphacelariales Migula 1908 . Chaetopteris , Halopteris , Stypocaulon , Sphacelaria , Verosphacella .
Sporochnales Sauvageau 1926 . Bellotia , Carpomitra , Nereia , Sporochonus , Tomaculopsis .
Syringodermatales Henry 1984 . Syringoderma .
Tilopteridales Bessey 1907 . Cutleria , Halosiphon , Haplospora , Phaeosiphoniella , Phyllaria , Tilopteris .
Phaeosacciophyceae R.A.Andersen, L.Graf & H.S.Yoon 2020 [ 59]
Phaeothamiophyceae Andersen & Bailey in Bailey et al. 1998
Schizocladiophyceae Henry, Okuda & Kawai 2003 . Schizocladia .
Xanthophyceae Allorge 1930, emend. Fritsch 1935 [Heterokontae Luther 1899 ; Heteromonadea Leedale 1983; Xanthophyta Hibberd 1990 ]
Diatomista Derelle et al. 2016, emend. Cavalier-Smith 2017 [SIII clade]. Incertae sedis : environmental clades MOCH-1, MOCH-2 and MOCH-4 Massana et al. 2014 .
Bolidophyceae Guillou et al. 1999 [Parmales Booth & Marchant 1987 ]. Bolidomonas , Triparma , Tetraparma , Pentalamina .
Diatomeae Dumortier 1821 [Bacillariophyta Haeckel 1878 ]
Leptocylindrophytina D.G. Mann, in Adl et al. 2019
Ellerbeckiophytina D.G. Mann, in Adl et al. 2019 . Ellerbeckia .
Probosciophytina D.G. Mann, in Adl et al. 2019 . Proboscia .
Melosirophytina D.G. Mann, in Adl et al. 2019 . Aulacoseira , Melosira , Hyalodiscus , Stephanopyxis , Paralia , Endictya .
Coscinodiscophytina Medlin & Kaczmarska 2004, emend. Adl et al. 2019 . Actinoptychus , Coscinodiscus , Actinocyclus , Asteromphalus , Aulacodiscus , Stellarima .
Rhizosoleniophytina D.G. Mann, in Adl et al. 2019 . Guinardia , Rhizosolenia , Pseudosolenia .
Arachnoidiscophytina D.G. Mann, in Adl et al. 2019 . Arachnoidiscus .
Bacillariophytina Medlin & Kaczmarska 2004, emend. Adl et al. 2019
Mediophyceae Jouse & Proshkina-Lavrenko in Medlin & Kaczmarska 2004
Chaetocerotophycidae Round & R.M. Crawford in Round et al. 1990, emend. Adl et al. 2019 . Chaetoceros , Bacteriastrum , Dactyliosolen , Cerataulina , Hemiaulus , Eucampia , Acanthoceras , Urosolenia , Terpsinoë , Hydrosera .
Lithodesmiophycidae Round & R.M. Crawford in Round et al. 1990, emend. Adl et al. 2019 . Lithodesmium , Lithodesmioides , Helicotheca , Bellerochea , Ditylum .
Thalassiosirophycidae Round & R.M. Crawford in Round et al. 1990 . Thalassiosira , Lindavia , Cyclotella , Stephanodiscus , Cyclostephanos , Discostella , Bacteriosira , Skeletonema , Detonula .
Cymatosirophycidae Round & R.M. Crawford in Round et al. 1990 . Cymatosira , Minutocellus , Papiliocellulus , Leyanella , Extubocellulus , Plagiogrammopsis , Campylosira , Brockmanniella , Pierrecomperia .
Odontellophycidae D. G. Mann, in Adl et al. 2019 . Odontella , Triceratium , Cerataulus , Pleurosira , Pseudauliscus , Amphitetras , Trieres , Mastodiscus .
Chrysanthemodiscophycidae D.G. Mann, in Adl et al. 2019 . Chrysanthemodiscus , Biddulphiopsis , Trigonium , Isthmia , Lampriscus , Stictocyclus , Ardissonea , Climacosphenia , Toxarium .
Biddulphiophyceae D. G. Mann, in Adl et al. 2019
Bacillariophyceae Haeckel 1878, emend. Adl et al. 2019
Striatellaceae Kützing 1844 . Striatella , Pseudostriatella .
Urneidophycidae Medlin 2016 . (P?) Plagiogramma , Dimeregramma , Rhaphoneis , Delphineis , Psammoneis , Bleakeleya , Asterionellopsis .
Fragilariophycidae Round in Round, Crawford & Mann 1990, emend. Adl et al. 2019 . Fragilaria , Synedra , Tabellaria , Asterionella , Diatoma , Tabularia , Cyclophora , Astrosyne , Licmophora , Rhabdonema , Grammatophora , Staurosira , Thalassionema .
Bacillariophycidae D.G. Mann in Round, Crawford & Mann 1990, emend. Adl et al. 2019 . Eunotia , Achnanthes , Bacillaria , Nitzschia , Pseudonitzschia , Cylindrotheca , Navicula , Seminavis , Haslea , Pleurosigma , Gyrosigma , Achnanthidium , Cocconeis , Frustulia , Diploneis , Sellaphora , Pinnularia , Gomphonema , Cymbella , Didymosphenia , Phaeodactylum , Amphora , Entomoneis , Epithemia , Surirella , etc.
Dictyochophyceae Silva 1980 [Silicoflagellata Lemmermann, 1901 ]
Dictyochales Haeckel 1894 [Dictyochida Borgert, 1891 ; Dictyochaceae Lemmermann, 1901 ]. Dictyocha .
Florenciellales Eikrem, Edvardsen & Throndsen 2007 .[ 61] Florenciella , Verrucophora .
Pedinellales Zimmermann et al. 1984 [Ciliophryida Febre-Chevalier, 1985 ; Actinomonadaceae Kent, 1880 ]. Actinomonas , Apedinella , Ciliophrys , Mesopedinella , Palatinella , Pedinella , Pseudopedinella , Pteridomonas .
Rhizochromulinales O'Kelly & Wujek 1994 [Rhizochromulinaceae O'Kelly & Wujek, 1994 ]. Rhizochromulina .
Pelagophyceae Andersen & Saunders 1993
Pelagomonadales Andersen & Saunders 1993 . Aureococcus , Pelagococcus , Pelagomonas .
Sarcinochrysidales Gayral & Billard 1977
Sarcinochrysidaceae Gayral & Billard 1977 . Ankylochrisis , Arachnochrysis , Aureoscheda , Aureoumbra , Nematochrysopsis , Pelagospilus , Pulvinaria , Sarcinochrysis , Sargassococcus .
Chrysocystaceae M.Melkonian, H.S.Yoon & R.A.Andersen 2018 .[ 62] Chrysocystis , Chrysoreinhardia , Sungminbooa .
Plocamiomonadales Daugbjerg & Lovejoy 2024 . Plocamiomonas .[ 63]
Cladogram of Alveolata , based on a SSU rDNA phylogeny of basal alveolates[ 64] and a 203-gene phylogeny of apicomplexans and their relatives[ 65] published in 2023.
Colponemida Cavalier-Smith 1993 emend. Adl et al. 2019 (P)
Colpodellida Cavalier-Smith 1993, emend. Adl et al. 2005, 2019 ["chrompodellids" Janouškovec et al. 2015 ;[ 66] Chromeridophyta R.B.Moore et al., 2008 ; Colpodellophyceae Molinari & Guiri 2023 ; Chromerida Guiry 2024 ].[ 67] Incertae sedis : Piridium .[ 68]
Perkinsozoa Norén & Moestrup in Norén et al 1999 [Perkinsea Levine 1978 ]. Incertae sedis genus: Phagodinium , Rastrimonas .[ p]
Maranthos Alacid & Reñé 2021 [ 70]
Pararosariidae Jeon & Park 2021 . Pararosarium .[ 71]
Parviluciferaceae Reñé & Alacid 2017 . Dinovorax , Parvilucifera , Snorkelia ,[ 72] Tuberlatum .[ 73]
Perkinsidae Levine 1978 . Perkinsus .
Xcellidae Freeman, Fuss, Kristmundsson, Keeling, Shalchian-Tabrizi & Bass 2017 em. Karlsbakk, Nystøyl, Plarre & Nylund 2021 . Cryoxcellia ,[ 74] Gadixcellia , Notoxcellia ,[ 75] Salmoxcellia ,[ 76] Xcellia .
Dinoflagellata Bütschli 1885, emend. Fensome et al. 1993, emend. Adl et al. 2005 . Incertae sedis genera: free-living (Adenoides , Akashiwo , Amphidiniella , Ankistrodinium , Apicoporus , Archaeosphaerodiniopsis , Bispinodinium , Bysmatrum , Cabra , Cladopyxis , Crypthecodinium , Cucumeridinium , Dactylodinium , Dicroerisma , Gloeodinium , Grammatodinium , Gynogonadinium , Gyrodiniellum , Halostylodinium , Heterodinium , Moestrupia , Paragymnodinium , Phytodinium , Plagiodinium , Planodinium , Pileidinium , Pseudadenoides , Pseudothecadinium , Pyramidodinium , Rhinodinium , Roscoffia , Sabulodinium , Sphaerodinium , Spiniferodinium , Testudodinium , Thecadinium , Thecadiniopsis , Togula ) and parasitic Blastodiniales Chatton 1906 [no longer valid] (Amyloodinium , Apodinium , Cachonella , Caryotoma , Crepidoodinium , Haplozoon , Oodinium , Myxodinium , Piscinoodinium , Protoodinium , Rhinodinium , Schyzochitriodinium , Stylodinium ).
Psammosa Okamoto, A. Horák & Keeling 2012 [ 77] [ 78]
Oxyrrhis Dujardin 1841 [Oxyrrhinaceae Sournia 1984 ][ 78]
Eleftheros Tikhonenkov, Hehenberger & Keeling 2023 [ 78]
Ichthyodinida Cavalier-Smith 2018 [environmental Marine Alveolate Group I][ 78]
Syndiniales Loeblich III 1976 . Incertae sedis : Ellobiopsidae (Ellobiopsis , Thalassomyces ).
Noctilucales Haeckel 1894 [Noctiluciphyceae Fensome et al. 1993 ]. Abedinium , Cachonodinium , Craspedotella , Cymbodinium , Kofoidinium , Leptodiscus , Noctiluca , Petalodinium , Pomatodinium , Scaphodinium , Spatulodinium .
Dinophyceae Pascher 1914
Gymnodiniphycidae Fensome et al. 1993
Gymnodiniales F. Stein 1878, emend. G. Hansen & Moestrup in Daugbjerg et al. 2000 sensu stricto . Barrufeta , Chytriodinium , Dissodinium , Erythropsidinium , Greuetodinium , Gymnodinium , Lepidodinium , Nematodinium , Nusuttodinium , Pellucidodinium , Polykrikos , Proterythropsis , Wangodinium ,[ 80] Warnowia .
Amphidinium Claparède & Lachmann 1859, emend. Flø Jørgensen et al. 2004 sensu stricto
Gyrodinium Kofoid & Swezy 1921, emend. G. Hansen & Moestrup in Daugbjerg et al. 2000 sensu stricto
Kareniaceae Bergholtz et al. 2005 . Brachidinium , Karenia , Karlodinium , Takayama .
Ceratoperidiniaceae Loeblich III 1980, emend. Reñé & de Salas 2013 . Ceratoperidinium , Kirithra .
Torodiniales Boutrup, Moestrup & Daugbjerg 2016 . Kapelodinium , Torodinium .
Levanderina Moestrup et al. 2014
Margalefidinium F. Gómez, Richlen & D.M. Anderson 2017
Cochlodinium F. Schütt emend. F. Gómez, Richlen & D.M. Anderson 2017 sensu stricto
Ptychodiscales Fensome et al. 1993 . Achradina , Amphilothus , Balechina , Ptychodiscus , Sclerodinium .
Borghiellaceae Moestrup, Lindberg, & Daugbjerg 2009 . Baldinia , Borghiella .
Tovelliaceae Moestrup et al. 2005 . Bernardinium , Esoptrodinium , Jadwigia , Tovellia .
Suessiaceae Fensome et al. 1993, emend. Moestrup et al. 2009 . Ansanella , Asulcocephalium , Biecheleria , Biecheleriopsis , Leiocephalium , Pelagodinium , Polarella , Prosoaulax , Protodinium , Symbiodinium , Yihiella .
Peridiniphycidae Fensome et al. 1993
Gonyaulacales Taylor 1980 . Alexandrium , Amylax , Ceratium , Ceratocorys , Coolia , Fukuyoa , Fragilidium , Gambierdiscus , Goniodoma , Gonyaulax , Lingulodinium , Ostreopsis , Pentaplacodinium , Peridiniella , Protoceratium , Pyrocystis , Pyrodinium , Pyrophacus , Tripos .
Peridiniales Haeckel 1894 . Amphidiniopsis , Archaeperidinium , Blastodinium *, Diplopelta , Diplopsalis , Diplopsalopsis , Herdmania , Niea , Oblea , Palatinus , Parvodinium , Peridinium , Peridiniopsis , Preperidinium , Protoperidinium , Qia , Vulcanodinium .
Thoracosphaeraceae Schiller 1930 . Aduncodinium , Amyloodinium , Apocalathium , Blastodinium , Chimonodinium , Cryptoperidiniopsis , Duboscquodinium , Ensiculifera , Laciniporus ,[ 81] Leonella , Luciella , Naiadinium , Paulsenella , Pentapharsodinium , Pfiesteria , Scrippsiella , Stoeckeria , Theleodinium , Thoracosphaera , Tintinnophagus , Tyrannodinium .
Podolampadaceae Lindemann 1928 . Blepharocysta , Gaarderiella , Heterobractum , Lissodinium , Mysticella , Podolampas .
Kryptoperidiniaceae Lindemann 1925 ["dinotoms"]. Blixaea , Durinskia , Galeidinium , Kryptoperidinium , Unruhdinium , etc.
Heterocapsaceae Fensome et al. 1993 . Heterocapsa .
Amphidomataceae Sournia 1984 . Amphidoma , Azadinium .
Oxytoxaceae Lindemann 1928 . Corythodinium , Oxytoxum .
Centrodiniaceae Hernández-Becerril 2010 . Centrodinium .
Dinophysales Kofoid 1926 . Amphisolenia , Citharistes , Dinofurcula , Dinophysis , Histioneis , Latifascia , Metadinophysis , Metaphalacroma , Ornithocercus , Oxyphysis , Parahistioneis , Phalacroma , Pseudophalacroma , Sinophysis , Triposolenia .
Prorocentrales Lemmermann 1910 . Mesoporus , Prorocentrum .
Squirmida Cavalier-Smith 2014 . Digyalum , Filipodium , Platyproteum .[ 82] [ 83]
Apicomplexa Levine 1980, emend. Adl et al. 2005 . Incertae sedis genera: Aggregata [ q] Frenzel 1885 , Christalloidophora Dehorne 1934 , Dobellia * Ikeda 1914 , Echinococcidium * Porchet 1978 , Globidiellum * Brumpt 1913 , Joyeuxella * Brasil 1902 , Rhabdospora * Laguesse 1906 , Spermatobium * Eisen 1895 , Spiriopsis * Arvy & Peters 1972 , Spirogregarina * Wood & Herman 1943 , Toxocystis * Léger & Duboscq 1910 , Trophosphaera * Le Calvez 1939 .
?Agamococcidiorida Levine 1979 . Gemmocystis *, Rhytidocystis .
?Protococcidiorida Kheisin 1956 . Angeiocystis *, Coelotropha *, Grellia *, Eleutheroschizon *, Mackinnonia *, Myriosporides *, Myriospora *, Sawayella *.
Aconoidasida Mehlhorn et al. 1980 [Hematozoa Vivier 1982 ] (P)
Haemospororida Danilewsky 1885 . Dionisia *, Haemocystidium , Haemoproteus , Hepatocystis , Leucocytozoon , Mesnilium *, Nycteria , Parahaemoproteus , Plasmodium , Polychromophilus , Rayella *, Saurocytozoon *.
Piroplasmorida Wenyon 1926 . Anthemosoma *, Babesia , Cytauxzoon , Echinozoon *, Haemohormidium *, Sauroplasma *, Serpentoplasma *, Theileria .
Nephromycida Cavalier-Smith 1993, emend. Adl et al. 2019 . Nephromyces , Cardiosporidium .
Conoidasida Levine 1988 (P)
Coccidia Leuckart 1879 (P)
Adeleorina Léger 1911 . Adelea *, Adelina , Babesiosima *, Bartazoon *, Chagasella *, Cyrilia *, Dactylosoma , Desseria *, Ganapatiella *, Gibbsia *, Haemogregarina , Haemolivia , Hepatozoon , Ithania *, Karyolysus , Klossia , Klossiella *, Legerella *, Orcheobius *, Rasajeyna *.
Eimeriorina Léger 1911 . Atoxoplasma , Barrouxia *, Besnoitia , Caryospora , Caryotropha , Choleoeimeria , Cyclospora , Cystoisospora , Defretinella *, Diaspora *, Dorisa *, Dorisiella *, Eimeria , Elleipsisoma *, Goussia , Hammondia , Hyaloklossia , Isospora , Lankesterella , Mantonella *, Merocystis , Neospora , Nephroisospora , Ovivora *, Pfeifferinella , Pseudoklossia , Sarcocystis , Schellackia , Selenococcidium *, Selysina *, Spirocystis *, Toxoplasma , Tyzzeria *, Wenyonella *.
Gregarinasina Dufour 1828 (P). Incertae sedis genera: Digyalum Koura et al. 1990 , Exoschizon Hukui 1939 .
Archigregarinorida Grassé 1953 (P). Meroselenidium , Merogregarina , Selenidium , Selenocystis , Veloxidium .
Eugregarinorida Léger 1900 (P). Apolocystis *, Amoebogregarina , Ancora , Ascogregarina , Asterophora , Blabericola , Caliculium , Cephaloidophora , Colepismatophila , Cystobia *, Cystocephalus *, Difficilina , Diplauxis *, Enterocystis , Ganymedes , Geneiorhynchus , Gonospora , Gregarina , Heliospora , Hentschelia *, Hirmocystis *, Hoplorhynchus , Lankesteria , Lecudina , Leidyana , Lithocystis , Monocystella *, Monocystis , Nematopsis , Nematocystis *, Neoasterophora *, Paralecudina , Paraschneideria , Phyllochaetopterus , Pileocephalus *, Polyplicarium , Polyrhabdina , Porospora *, Prismatospora , Protomagalhaensia , Psychodiella , Pterospora , Pyxinia , Pyxinioides , Rhabdocystis *, Sphaerorhynchus *, Steinina , Stenophora , Stylocephalus , Sycia *, Syncystis , Thalicola *, Thiriotia , Trichotokara , Uradiophora *, Urospora , Xiphocephalus , Zygosoma *, and more.
Neogregarinorida Grassé 1953 . Apicystis , Aranciocystis , Caulleryella *, Coelogregarina *, Farinocystis *, Gigaductus *, Lipocystis *, Lipotropha *, Lymphotropha *, Machadoella *, Mattesia , Menzbieria , Ophryocystis , Schizocystis *, Syncystis , Tipulocystis *.
Cryptogregarinorida Cavalier-Smith 2014, emend. Adl et al. 2019 . Cryptosporidium .
Blastogregarinea Chatton and Villeneuve 1936, emend. Simdyanov et al. 2018 . Chattonaria , Siedleckia .
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
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)
Oligotrichea Bütschli 1887
Licnophoridae Bütschli 1887 . Licnophora , Prolicnophora .
Kiitrichidae Nozawa 1941 . Caryotricha , Kiitricha .
Lamellicorticata Vd'ačný et al. 2010
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
Colpodea Small & Lynn 1981
Nassophorea (P)
Prostomatea Schewiakoff 1896 (P)
Apsiktratidae Foissner et al. 1994 [Prostomatida Schewiakoff 1896 ]
Plagiopylea Small & Lynn 1985
Oligohymenophorea de Puytorac et al. 1974
Apostomatia Chatton & Lwoff 1928
Astomatia Schewiakoff 1896
Hymenostomatia Delage & Hérouard 1896
Peniculia Fauré-Fremiet in Corliss 1956 (P)
Peniculida Fauré-Fremiet in Corliss 1956
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
Scuticociliatia Small 1967 (P)
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 [ edit ]
Cercozoa incertae sedis :
Aquavolonida Bass & Berney 2018 . Aquavolon , Lapot .[ 86]
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)
Cercomonadida [ h] Poche 1913, emend. Vickerman 1983, emend. Mylnikov 1986, emend. Karpov et al. 2006; emend Howe et al. 2009; emend Cavalier-Smith 2012 [Cercomonadidae Kent 1880, emend. Mylnikov and Karpov 2004 ; Cercobodonidae Hollande 1942 ]. Cercomonas , Eoercomonas , Filomonas , Neocercomonas , Cavernomonas .
Paracercomonadida Cavalier-Smith 2018 [Paracercomonadidae Cavalier-Smith & Karpov 2012 ]. Brevimastigomonas , Metabolomonas , Nucleocercomonas , Paracercomonas , Phytocercomonas .
Glissomonadida Howe & Cavalier-Smith 2009 [Heteromitidae Kent 1880, emend. Mylnikov 1990, emend. Mylnikov & Karpov 2004 ]
Saccharomycomorphidae Feng, He, Jiang, Zhang & Yu 2021 .[ 87] Saccharomycomorpha .
Sandonidae Howe et al. 2009 . Sandona , Flectomonas , Neoheteromita , Mollimonas .
Dujardinidae Howe & Cavalier-Smith 2011 . Dujardina .
Bodomorphidae Hollande 1952, emend. Cavalier-Smith in Howe et al. 2009 . Bodomorpha .
Proleptomonadidae Howe et al. 2009 . Proleptomonas .
Allapsidae Howe & Cavalier-Smith 2009 . Allapsa , Teretomonas , Allantion .
Viridiraptoridae Hess & Melkonian 2013 . Orciraptor , Viridiraptor .
Pansomonadidae Vickerman in Vickerman et al. 2005 . Agitata , Aurigamonas .
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
Ventricleftida Cavalier-Smith 2011
Tectofilosida Cavalier-Smith 2003
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
Variglissida Cavalier-Smith 2014
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 .[ 89]
Cyphoderiidae de Saedeleer 1934 . Campascus *, Corythionella , Cyphoderia [Schaudinnula ], Messemvriella *, Pseudocorythion , Ichthyosquama , Knarr , Oleiformis , Psammoderia .[ 90]
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 .[ 91] Leptogromia , Trivalvularis .
Metromonadea Cavalier-Smith 2007, emend. Cavalier-Smith 2011 . Metromonas , Metopion , Micrometopion , Kiitoksia .
Granofilosea Cavalier-Smith & Bass 2009 .[ 92] 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 [ edit ]
Vampyrellida West 1901, emend. Hess et al. 2012 [Aconchulinida De Saedeleer 1934 ]. Incertae sedis : Arachnula , Gobiella , Lateromyxa , Penardia .[ 93]
Phytomyxea Engler & Prantl 1897
Marinomyxa Kolátková, Čepička, Hoffman & Vohník 2021 [ 95]
Plasmodiophorida Cook 1928 [Plasmodiophorales Engl. 1892 ;[ 96] Plasmodiophoromycota R.H. Whittaker 1969 [ 1] ]. Plasmodiophora [Ostenfeldiella ], Polymyxa , Woronina , Ligniera [Anisomyxa ; Rhizomyxa ; Sorolpidium ], Sorosphaerula [Tuburcinia ; Sorosporium ], Spongospora [Clathrosorus ], ?Membranosorus , ?Octomyxa , ?Polymyxa , ?Sorodiscus , ?Tetramyxa [Molliardia ; Thecaphora ].[ 97] [ 98] [ 99]
Phagomyxida Cavalier-Smith 1993 . Phagomyxa , Maullinia .
Filoreta Bass & Cavalier-Smith 2009
Gromia Dujardin 1835
Ascetosporea Sprague 1979, emend. Cavalier-Smith 2009
Foraminifera d'Orbigny 1826 . Incertae sedis groups: ?Lagenida , ?Heterogromia , ?Komokiacea *.
Monothalamids Pawlowski et al. 2013 (P) [currently under revision].[ 100] All genera traditionally in Allogromiida , Astrorhizida , and the Xenophyophorea . Genera unplaced into families: Allogromia , Astrammina , Crithionina , Notodendrodes , Psammophaga , Bathysiphon .
Tubothalamea Pawlowski et al. 2012 . † Fusulinida Delage & Hérouard 1896 probably belongs to this group.[ 3]
Globothalamea Pawlowski et al. 2012
Rotaliida Delage & Hérouard 1896
Globigerinida Lankester 1885 emend. Arz, Arenillas & Gilabert 2022 [ 7] (P)
† Parvularugoglobigerinoidea Arenillas, Arz & Gilabert 2022
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
† Hantkeninoidea Cushman 1927
† 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
† Favuselloidea Longoria 1974
† 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
† 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]
Order ? (unnamed)[ 7]
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 [ 106]
† Entactinaria Kozur & Mostler 1982 sensu Nakamura & Suzuki 2020 [ t]
† Entactiniidae Riedel 1967 . Afanasievella , Apophysiactinia , Borisella , Callela , Cancellosphaera , Costaentactinia , Cyclocarpus nomen dubium , Duodecimentactinia , Langsonsphaera ,[ 107] 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 .[ 108]
† Tetrentactiniidae Kozur & Mostler 1979 . Multisphaera .
† Triassothamnidae Takahashi, Maekawa & Dumitrica 2022 . Triassothamnus .[ 107]
Haptophyta Hibberd 1976, ex. Edvardsen & Eikrem 2000
Pavlovales Green 1976 [Pavlovophyceae Cavalier-Smith 1986 ]. Diacronema , Exanthemachrysis , Pavlova , Rebecca .
Prymnesiophyceae Hibberd 1976
Prymnesiales Papenfuss 1955, emend. Edvardsen & Eikrem 2000 . Chrysochromulina , Chrysocampanula , Dicrateria , Haptolina , Prymnesium , Pseudohaptolina .
Phaeocystales Medlin 2000 . Phaeocystis .
Isochrysidales Pascher 1910, emend. Edvardsen & Eikrem 2000 . Emiliania , Gephyrocapsa , Isochrysis , Ruttnera , Tisochrysis .
Coccolithales Schwarz 1932, emend. Edvardsen & Eikrem 2000 . Balaniger , Calciosolenia , Coccolithus , Hymenomonas , Chrysotila , Wigwamma .
Rappephyceae M. Kawachi, R. Kamikawa & T. Nakayama 2021 [ 109]
Centroplasthelida Febvre-Chevalier & Febvre 1984 [Centrohelea Kühn 1926 sensu Cavalier-Smith in Yabuki et al. 2012 ; Centroheliozoa Dürrschmidt & Patterson 1987 ]
Cryptista Cavalier-Smith 1989, 2018, emend. Adl et al. 2019 [ edit ]
Barthelona Bernard, Simpson & Patterson 2000 [ 116]
Skoliomonas Eglit & Simpson 2024 [ 117]
Anaeramoebae [ 118] [Anaeramoebidae Táborský, Pánek & Čepička 2017 ]. Anaeramoeba .[ 119]
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
Oxymonadida Grassé 1952 . Barroella , Blattamonas , Brachymonas , Dinenympha , Microrhopalodina , Monocercomonoides , Notila , Opisthomitus , Oxymonas , Paranotila , Polymastix , Pyrsonympha , Sauromonas , Saccinobaculus , Streblomastix , Tubulimonoides .
Trimastigidae Saville Kent 1880-1882 . Trimastix .
Paratrimastigidae Zhang et al. 2015 . Paratrimastix .
Pharyngomonada Cavalier-Smith 2008 . Pharyngomonas
Tetramitia Cavalier-Smith 1993
Selenaionidae Hanousková et al. 2018 . Dactylomonas , Selenaion .
Neovahlkampfiidae Hanousková et al. 2018 . Neovahlkampfia .
Eutetramitia Hanousková et al. 2018
Vahlkampfiidae Jollos 1917 (P). Fumarolamoeba , Heteramoeba , Naegleria , Neovahlkampfia , Paravahlkampfia , Tetramitus , Vahlkampfia , Willaertia .
Gruberellidae Page & Blanton 1985 . Gruberella *, Stachyamoeba .
Acrasidae Poche 1913 . Acrasis , Allovahlkampfia , Pocheina .
Percolomonadidae Cavalier-Smith 2008 . Percolomonas .
Psalteriomonadidae Cavalier-Smith 1993 . Harpagon , Psalteriomonas , Monopylocystis , Pseudoharpagon , Sawyeria .
Stephanopogonidae Corliss 1961 . Stephanopogon .
Creneidae Pánek et al. 2014 . Creneis .
Tulamoebidae Kirby et al. 2015 . Pleurostomum , Tulamoeba .
Euglenida Butschli 1884, emend. Simpson 1997 . Uncertain genera: † Moyeria Thusu 1973 .[ 123]
Petalomonadida Cavalier-Smith 1993 . Atraktomonas *, Biundula *, Calycimonas , Dolium , Dylakosoma *, Notosolenus , Pentamonas *, Petalomonas , Scytomonas , Sphenomonas , Tropidoscyphus *.[ 124]
Alistosa Lax & Simpson 2021 . Ploeotia , Serpenomonas , Keelungia , Lentomonas , Decastava .[ 125]
Entosiphon Cavalier-Smith, Chao & Vickerman 2016 [ 126] [ 127]
Gaulosia Lax, Cho & Keeling 2023 [ 127]
Karavia Lax, Cho & Keeling 2023 . Hemiolia , Liburna .[ 127]
Olkaspira Lax & Simpson 2021 .[ 125]
Chelandium Lax, Cho & Keeling 2023 [ 127]
Olkasia Lax, Lee, Eglit & Simpson 2019 [ 128]
Spirocuta Cavalier-Smith, Chao & Vickerman 2016 [ 126] [Helicales Perschke et al. 2017 ][ 125]
Anisonemia Cavalier-Smith 2016
Peranemidae Bütschli 1884 (P). Chasmostoma , Jenningsia [Peranemopsis ],[ 130] Peranema , Teloprocta , Urceolus [Urceolopsis , Phialonema ].[ 131] [ 129] [ 125] [ 132]
Euglenophyceae Schoenichen 1925, emend. Marin & Melkonian 2003 [Euglenea Butschli 1884, emend. Busse & Preisfeld 2002 ]
Rapaza Yamaguchi et al. 2012
Eutreptiales Leedale 1967, emend. Marin & Melkonian 2003 . Eutreptia , Eutreptiella .
Euglenales Leedale 1967, emend. Marin & Melkonian 2003
Phacaceae Kim et al. 2010 . Discoplastis , Lepocinclis , Phacus .
Euglenaceae Dujardin 1841, emend. Kim et al. 2010 . Ascoglena , Colacium , Cryptoglena , Euglena , Euglenaformis , Euglenamorpha , Euglenaria , Euglenopsis , Hegneria , Klebsina , Monomorphina , Strombomonas , Trachelomonas .
Diplonemea Cavalier-Smith 1993, emend. Simpson 1997
Symbiontida Yubuki et al. 2009 . Bihospites , Calkinsia , Postgaardi .
Kinetoplastea Honigberg 1963 . Incertae sedis genera: ?Bordnamonas , ?Cephalothamnium , ?Rhynchoidomonas .
Prokinetoplastina Vickerman in Moreira et al. 2004 . Ichthyobodo , Perkinsela .
Metakinetoplastina Vickerman in Moreira et al. 2004
Neobodonida Vickerman in Moreira et al. 2004 . Actuariola , Avlakibodo ,[ 133] Azumiobodo , Cruzella , Cryptaulaxella ,[ 134] Dimastigella , Klosteria , Neobodo , Phanerobia , Rhynchobodo , Rhynchomonas .
Parabodonida Vickerman in Moreira et al. 2004 . Cryptobia , Jarrellia , Parabodo , Procryptobia , Trypanoplasma .
Eubodonida Vickerman in Moreira et al. 2004 . Bodo .
Trypanosomatida Kent 1880, emend. Vickerman in Moreira et al. 2004 . Angomonas , Blechomonas , Leptomonas , Paratrypanosoma , Sergeia , Strigomonas , Wallaceina , Phytomonas , Trypanosoma .
Diphyllatea Cavalier-Smith 2003 → Diphylleida Cavalier-Smith 1993 [Anisomonadea Cavalier-Smith 1993 emend. 2000 ] → Collodictyonidae Brugerolle et al. 2002, emend. Adl et al. 2019 [Diphylleidae Cavalier-Smith 1993 ; Sulcomonadidae Cavalier-Smith 2013 ]: Collodictyon Carter 1865 , Diphylleia Massart 1920 [Aulacomonas Skuja 1939 ], Sulcomonas Brugerolle 2006 .
Hilomonadea Cavalier-Smith 2008 emend. 2012 → Rigifilida Cavalier-Smith in Yabuki et al. 2012 [Micronucleariida Cavalier-Smith 2008 ]
Genera of uncertain affiliation [ edit ]
Acinetactis , Actinastrum , Actinocoma , Actinolophus , Adinomonas , Aletium , Amphimonas , Amylophagus , Aphelidiopsis , Asterocaelum , Asthmatos , Aurospora , Barbetia , Berkeleyaesol ,[ 137] Belaria , Belonocystis , Bertarellia , Bertramia , Bodopsis , Boekelovia , Branchipocola , Camptoptyche , Chalarodora , Cibdelia , Cichkovia , Cinetidomyxa , Cingula , Cladomonas , Clathrella , Codonoeca , Coelosporidium [ v] , 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 ,[ 138] Gymnococcus , Gymnophrydium , Haematotractidium , Hartmannina , Heliobodo , Heliomonas , Hermisenella , Heterogromia , Hillea , Hyalodaktylethra , Immanoplasma , Isoselmis , Janickina , Kamera , Lagenidiopsis , Liegeosia , Luffisphaera [ w] , Lymphocytozoon , Lymphosporidium , Macappella , Magosphaera , Malpighiella , Martineziella , Megamoebomyxa , Meringosphaera , Microcometes , Monochrysis , Monodus , Mononema , Myrmicisporidium , Naupliicola , Nephrodinium , Neurosporidium , Orbulinella , Ovicola , Palisporomonas , Pansporella , Paradinemula , Parakaryon [ x] , 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 .[ 3]
^ The first eukaryotes were “neither plants, animals, nor fungi”, hence as defined, the Protista would include the earliest common ancestor of all eukaryotes.
^ In the original 4-kingdom model proposed in 1959, Protista included all unicellular microorganisms such as bacteria .
^ a b This taxon belongs to the paraphyletic phylum Apusozoa Cavalier-Smith 1997 emend. 2013 .[ 15]
^ 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]
^ 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]
^ 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]
^ 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]
^ 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.[ 23] [ 3]
^ This genus groups with Dermamoebidae in a phylogenetic anaysis that uses a limited number of taxa.[ 32] In the recent taxonomy it is listed as a separate clade until its position is better resolved.[ 3]
^ This genus could be a synonym of Vannella .[ 3]
^ 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]
^ Latest analyses show that Microheliella is the sister group to Cryptista .[ 12]
^ Telonemia is the sister group to the SAR clade.[ 9]
^ The picobiliphytes are phylogenetically closer to Rhodophyta and Rhodelphis than to other groups, and are therefore considered part of Archaeplastida .[ 38]
^ Cavalier-Smith and Scoble (2013) show a sister relationship between raphidophytes and Actinophryida .[ 57]
^ Apart from morphological descriptions, no molecular analysis has solidly affiliated Rastrimonas with the Perkinsozoa .[ 69]
^ Highly divergent 18S rRNA.[ 3]
^ This taxon is artificial; many of the families listed in it are not monophyletic and have little support from phylogenetic analyses.[ 3]
^ There is only one molecular sequence from this genus, which causes long branches in gene phylogenies , and the node is unresolved.[ 3]
^ 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.[ 106]
^ The genus Tetragonidium is known only from one written diagnosis and some illustrations, and its affinities with Cryptophyceae are very uncertain.[ 114]
^ Probably a synonym of the zygomycete fungus Nephridiophaga .[ 3]
^ May be the same genus as Belonocystis .[ 3]
^ It is not yet known whether this organism is an eukaryote or a prokaryote.[ 139]
^ a b Whittaker, R. H. (1969). "New Concepts of Kingdoms of Organisms" . Science . 163 (3863): 150–160. Bibcode :1969Sci...163..150W . doi :10.1126/SCIENCE.163.3863.150 . PMID 5762760 .
^ Ruggiero, Michael A.; Gordon, Dennis P.; Orrell, Thomas M.; Bailly, Nicolas; Bourgoin, Thierry; Brusca, Richard C.; Cavalier-Smith, Thomas; Guiry, Michael D.; Kirk, Paul M. (2015). "A Higher Level Classification of All Living Organisms" . PLOS ONE . 10 (4): e0119248. Bibcode :2015PLoSO..1019248R . doi :10.1371/JOURNAL.PONE.0119248 . PMC 4418965 . PMID 25923521 . (Erratum: doi :10.1371/journal.pone.0130114 )
^ a b c d e f g h i j k l m n o p q Adl, Sina M.; et al. (2019). "Revisions to the Classification, Nomenclature, and Diversity of Eukaryotes" . Journal of Eukaryotic Microbiology . 66 (1): 4–119. doi :10.1111/JEU.12691 . PMC 6492006 . PMID 30257078 .
^ a b c d e f g h i j k Leontyev, Dmitry V.; Schnittler, Martin; Stephenson, Steven L.; Novozhilov, Yuri K.; Shchepin, Oleg N. (2019). "Towards a phylogenetic classification of the Myxomycetes" . Phytotaxa . 399 (3): 209. doi :10.11646/PHYTOTAXA.399.3.5 .
^ a b Figueroa-Martinez F, Jackson C, Reyes-Prieto R (January 2019). "Plastid Genomes from Diverse Glaucophyte Genera Reveal a Largely Conserved Gene Content and Limited Architectural Diversity" . Genome Biology and Evolution . 11 (1): 174–188. doi :10.1093/gbe/evy268 . PMC 6330054 . PMID 30534986 .
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