Ceratiomyxomycetes綱(Ceratiomyxea) D. Hawksw., B. Sutton & Ainsw. 2019 , [ 35 ] Protosporangiida目Shadwick & Spiegel in Adl et al. 2012 (Ceratiomyxales GW Martin ex ML Farr & Alexop. )
Genera not assigned to any lower taxon: NeocolponemaGigeroff et al. 2023, LoeffelaGigeroff et al. 2023.[188]
Phylum[n]ColponemidiaTikhonenkov et al. 2014 → class ColponemeaCavalier-Smith 1993, emend. Tikhonenkov et al. 2014 → order ColponemidaCavalier-Smith 1993 → family ColponemidaeCavalier-Smith & Chao 2004, genus ColponemaStein 1878.
Rhizaria is a clade that contains a wide variety of primarily amoeboid protists with fine (filose) pseudopodia. It is generally divided into two phyla: Cercozoa, which is composed of a heterogeneous mixture of free-living flagellates, testate amoebae, obligate intracellular parasites, and photosynthetic amoebae, among others; and Retaria, which includes the radiolarians and forams, two groups that have a rich and continuous fossil record over 500 million years long. Discovered only through molecular phylogenetics,[218] Rhizaria is home to over 11,000 described species.[219]
Cercozoa
Cercozoa is a phylum of protists with extreme morphological diversity, ranging from small flagellates and amoeboflagellates, to testate amoebae, heliozoan- and radiolarian-like amoebae, and massive biomineralizing predators, as well as a few photosynthetic species. They are present abundantly in aquatic habitats and soils.[220] Cercozoa contains two subphyla: Filosa, represented primarily by free-living lineages, and Endomyxa, which includes obligate intracellular parasites and free-living predators.[221][s] Included in or related to Endomyxa are several clades known primarily from environmental sequences ('novel clades' 10, 11, 12),[222] with only a few members (grouped as Skiomonadea) formally described.[220]
The classification below is based largely on the 2019 revision by Adl et al.[6] and Cavalier-Smith et al.[8] Adl et al. excludes Endomyxa and Skiomonadea from Cercozoa, but later works generally include them.[221][220]
Phylum CercozoaCavalier-Smith 1998, emend. Adl et al. 2005
Order Glissomonadida(P)[230]Howe & Cavalier-Smith in Howe et al. 2009 (Heteromitidae Kent 1880, emend. Mylnikov 1990, emend. Mylnikov & Karpov 2004). Genus not assigned to any family: Neuromorpha.[230]
Superorder DiscomonadaCavalier-Smith in Cavalier-Smith, Chao & Lewis 2018, order DiscomonadidaCavalier-Smith in Scoble & Cavalier-Smith 2014, family DiscomonadidaeCavalier-Smith in Scoble & Cavalier-Smith 2014, genus Discomonas.
Order ReticulosidaCavalier-Smith in Cavalier-Smith & Chao 2003 emend. Bass et al. 2009, family FiloretidaeCavalier-Smith & Bass in Bass et al. 2009, genus Filoreta.
The foraminifera, or forams, are a diverse group of single-celled aquatic amoebae, usually bearing an organic or calcareous wall, amounting to over 6,700 living species and many fossil taxa.[257] They were originally classified as an order of protozoa (Foraminiferida) belonging to the now obsolete Granoreticulosa, along with other reticulose amoebae. Later they were separated into their own phylum by most protozoologists, although maintained as a class by micropaleontologists,[258] until they were placed as a subphylum of phylum Retaria along with the Radiolaria.[3] They are divided into three classes (or subclasses, when globally treated as a class): the paraphyletic monothalamids,[259] containing amoebae with single-chambered organic walls and a few naked amoebae (e.g., the large xenophyophores); and the monophyletic Globothalamea and Tubothalamea, which evolved from monothalamids, each containing amoebae with multi-chambered tests, either organic or calcareous.[256] Some alternate systems of dividing foraminifera into classes have been proposed,[260] but none have gained consensus. The relationships of monothalamids remain a work in progress due to the limited amount of genetic data.[256]
Foraminiferal classification was compiled numerous times through the 20th century, culminating in 1987 with the catalogue of foraminiferal genera published by paleontologists Alfred R. Loeblich Jr and Helen Tappan.[261] Later, catalogues have been published individually for separate groups of foraminifera on the basis of morphology and ecology. Two classifications for exclusively agglutinated (organic-walled) foraminiferal genera were published in 2001[258] and 2014,[262] and one classification for planktonic foraminifera was published in 2022.[263]
Disparia is a clade that unites several predatory protists, namely the provorans, hemimastigotes, and two mysterious genera Meteora and Solarion. It was described in 2025 through phylogenomic analyses that accompanied the discovery of Solarion.[285] Prior to these analyses, provorans, hemimastigotes and Meteora were considered 'orphan' taxa, with unstable phylogenetic positions. The first provoran, Ancoracysta twista, was briefly proposed to be related to Haptista.[8] However, in 2024 there was already evidence of a close affinity between Meteora, hemimastigotes, and provorans.[286] The phylum Caelestes, described in 2025, encompasses two monotypic genera of protists with a unique stalked extrusome used for immobilizing prey bacteria. Their morphology resembles that of celestial bodies, after which they are named. Only one of the genera, Solarion, is assigned to a family, and no classes or orders have been defined.[285] Hemimastigotes are a small group composed of ten species of heterotrophs with a pellicle and numerous flagella. First identified in 1988,[287] they remained an orphan group for decades due to the absence of genetic information, with taxonomists hypothesizing affinities to many different protists.[288] In 2018, two hemimastigotes were successfully sequenced and a phylogenomic analysis revealed their deep-branching placement near Diaphoretickes.[289] Between 2024 and 2025, a closer relationship with the clades Provora and Caelestes was resolved.[286][285] Provorans compose two ancient phyla of predatory microbes and a total of eight described species. They are small, fast-swimming predators capable of consuming larger prey, but are easily confused with other protists due to their unremarkable appearance; they required meticulous study to be recognized.[290][291][292]
The metamonads were first described by Pierre-Paul Grassé in the first volume of Traité de Zoologie, published in 1952, as the superorder Metamonadina or Anaxostylaria. They were originally composed of zooflagellates with four or more flagella, known as polymastigotes and hypermastigotes (e.g., Trichomonas, Oxymonas).[295] These were later split into other groups such as the Parabasalia and Preaxostyla (over 260 and 140 species respectively),[219][296] regarded as individual phyla. After rRNA phylogenetic analyses demonstrated their relatedness, the concept of metamonads was reintroduced by Cavalier-Smith as a monophyletic phylum Metamonada that includes both groups, as well as the anaerobic Fornicata (e.g., the free-living Carpediemonas and the parasitic Giardia;[297][298] around 140 species).[219] In this system, Preaxostyla is known as subphylum Anaeromonada, and Fornicata and Parabasalia are two infraphyla that belong to the subphylum Trichozoa.[4] Still, some taxonomists retain the Parabasalia at a phylum level.[299]
The classifications of Preaxostyla and Fornicata were last revised separately in the 2017 Handbook of the protists[300][301][302] and the higher classification of Metamonada was revised by Cavalier-Smith in 2021.[4] The classification of Parabasalia was completely updated in 2024.[299] Two new smaller clades of metamonads have been described in addition: anaeramoebae[303] and the 'BaSk' clade, containing barthelonids and skoliomonads.[304] However, Parabasalia was treated as a separate phylum by the 2024 revision, instead of a member of phylum Metamonada.[299]
Phylum MetamonadaGrassé 1952 emend.stat. n. Cavalier-Smith 2003
The phylum Heterolobosea contains around 170 species of amoebae, flagellates, and amoeboflagellates. It was initially established to unite two historically well-known amoeboid orders, Schizopyrenida (such as Naegleria fowleri, a human pathogen) and Acrasida (slime molds). Later, as more flagellates joined this grouping, the usage of Heterolobosea was split between two meanings: the more common usage applies to the entire clade,[6] while the usage by Cavalier-Smith and collaborators was restricted to a paraphyletic class of "traditional" heteroloboseans, with the name Percolozoa used for the phylum instead.[322][323] The first comprehensive phylogenomic study of Heterolobosea was published in 2025 by Tomáš Pánek and coauthors, resulting in its modern classification.[320]
Phylum HeteroloboseaPage & Blanton 1985 sensu Hanousková et al. 2019 (=Percolozoa Cavalier-Smith 1991)
The phylum Euglenozoa is home to at least 2,000 described species of single-celled flagellates of very dissimilar lifestyles. It was originally proposed to group the euglenids (such as the photosynthetic Euglena) and the kinetoplastids (like the pathogenic Trypanosoma), usually studied separatedly; eventually it included diplonemids and symbiontids as well.[328][329][330] Due to its share of photosynthetic species, traditionally regarded as algae, the phylum is also known as Euglenophyta by phycologists, and euglenids in particular were often studied as algae. Euglenids and kinetoplastids are the most diverse in terms of described species, although diplonemids may compose over 67,000 potential species.[330] The classification of euglenozoans was summarized by Cavalier-Smith in 2016.[329] A more phylogenetically precise revision of their classification was published in 2021 by Alexei Kostygov and coauthors,[330] with newer clades and genera described in the following years, particularly of euglenids. Still, one group of euglenids, the paraphyletic "ploeotids",[331] remains unresolved in the current classification, spread out across multiple clades.[332][333][334]
The phylum Telonemia (telonemids) contains a few species of flagellates found in ocean and fresh waters worldwide. It was originally proposed in 2006 for Telonema, a genus of previously uncertain affinity.[348] Under the Cavalier-Smith system, telonemids were initially classified as a class of Cryptista,[138] but later analyses consistently recovered it as a separate group.[25] Until 2019, only two species had been formally described, each belonging to a separate genus,[349] but environmental DNA sequencing suggested there were many more species not yet described.[350] Between 2022 and 2025, eight additional species were described along with three new genera, bringing the total number of species to ten.[351][352][353]
↑The first eukaryotes were "neither plants, animals, nor fungi", hence as defined, the Protista would include the earliest common ancestor of all eukaryotes.
↑The position of the genera Microcorycia, Parmulina, Penardochlamys and Zonomyxa, which were listed in 2002 under family Microcoryciidae, is not clear. They are placed here by morphological characters but this needs to be supported by molecular data.[6]
↑The 2019 revision by the ISOP ignores the grouping of some variosean genera into higher rank clades (orders and families) proposed in older studies, due to the weakly supported SSU rRNA phylogenetic analyses.[6]
12According to a 2018 molecular study: "The genera Coriophyllum, Neohalosacciocolax and Pseudorhododiscus are currently assigned to the Palmariales but have yet to be included in a molecular phylogenetic context, and their associations within the Palmariales remain equivocal."[82]
↑The 2019 revision by the ISOP inaccurately stated that goniomonads are classified as "Cyathomonadacea Pringsheim 1944".[6] The name Cyathomonadacea does not exist; Pringsheim only described the family Cyathomonadaceae based on the genus Cyathomonas, and later the order Cyathomonadales and class Cyathomonadea were described, each by different authors. However, as pointed out in 1993 by Gianfranco Novarino and Ian Lucas, this genus was based on a species that actually belonged to Goniomonas; since then, taxonomists have prioritized higher taxa named after it (Goniomonadaceae, Goniomonadales/adida, Goniomonadea) instead.[139]
↑The genus Tetragonidium is known only from one written diagnosis and some illustrations, and its affinities with cryptomonads are very uncertain.[139]
↑Within the order Anoecida, the four families Anoecaceae, Caecitellaceae, Cafeteriaceae and Symbiomonadaceae were included by T. Cavalier-Smith in 2006. In a 2013 revision, he simplified the classification by transferring Symbiomonadaceae and Anoecaceae to Cafeteriaceae.[7] However, both of those families remain accepted by the scientific community as independent from Cafeteriaceae as of 2020.[156]
↑O'Kelly placed the genera Antarctosaccion, Chrysomeris, Chrysonephos, Nematochrysis/Chrysowaernella, Phaeosaccion and Rhamnochrysis in the order Chrysomeridales nomen nudum in 1989,[174] and Cavalier-Smith placed it inside the class Chrysomeridophyceae (spelt originally as Chrysomerophycea) in 1995.[175] However, the class was proven to be polyphyletic. As a result, some of its genera (Nematochrysis/Chrysowaerella) were transferred to the class Chrysoparadoxophyceae, while others (Antarctosaccion and Phaeosaccion) were transferred to the class Phaeosacciophyceae. Due to a lack of molecular data, the placement of Chrysomeris, Chrysonephos and Rhamnochrysis remains unknown.[176]
↑The position of Eustigmatophyceae is still unstable, with different phylogenetic methodologies showing affinities to either of the SI and SII clades.[178][179]
12The taxonomic ranks above class level (phylum, subphylum, infraphylum, etc.) are unstable within the Alveolata, due to the many different existing classifications. For example, the phylum-level rank has been traditionally assigned to Perkinsozoa,[190] Dinoflagellata, Chromerida and Apicomplexa, all of which have been classified as phylum Myzozoa,[191] which in turn has been classified inside phylum Miozoa.[182] Treating the four myzozoan groups as separate phyla is generally accepted among phycologists,[192] while at the same time the rejection of higher taxonomic ranks due to being superfluous is prevalent among protistologists.[6]
↑Highly divergent 18S rRNA.[6] Since 2020 member of newly identified major apicomplexan subgroup Marosporida, putting together Aggregata octopianaFrenzel 1885, Merocystis kathaeDakin, 1911 (both Aggregatidae, originally coccidians), Rhytidocystis sp. 1 and Rhytidocystis sp. 2 Janouškovec et al. 2019 (RhytidocystidaeLevine, 1979, originally coccidians, Agamococcidiorida), and Margolisiella islandicaKristmundsson et al. 2011 (closely related to Rhytidocystidae)[194]
↑Apart from morphological descriptions, no molecular analysis has solidly affiliated Rastrimonas with the Perkinsozoa.[201]
↑The genera Parallobiopsis, Ellobiocystis and Rhizellobiopsis are only provisionally placed among ellobiopsids.[212]
↑This taxon is artificial; many of the families listed in it are not monophyletic and have little support from phylogenetic analyses.[6]
↑Whether Endomyxa is part of Cercozoa or not has been difficult to assess due to poor taxon sampling, but the most recent phylogenomic analyses support the inclusion of Endomyxa within Cercozoa.[217][220]
↑The family Massisteriidae was originally placed in the order Leucodictyida, along with the family Leucodictyidae which included two genera Leucodictyon and Reticulamoeba. However, Leucodictyon has not been genetically sequenced[225] and Reticulamoeba is not closely related to Massisteriidae.[226]
↑The monotypic genus[233]Nudifila branches within Pediglissa in the 2026 phylogenomic analysis.[220]
↑A proposed suborder-level clade by Cavalier-Smith in 2018 named Fiscullina[8] is not monophyletic because of uncultured tectofilosids.[236]
↑Regarding the classification of Retortamonadida, there is a disconnect between the Cavalier-Smith system and the system described by different authors in the Handbook of the protists and other studies. The order Retortamonadida was originally defined as two genera: Chilomastix and Retortamonas. In 2013, Cavalier-Smith modified the retortamonads to only include Retortamonas, and describing a separate order Chilomastigida for Chilomastix only, placed among the early branching Carpediemonas-like organisms.[309] However, this has not yet been recognized by later studies from the other authors.[302][310][311]
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12Cavalier-Smith, Thomas; Chao, Ema E.; Lewis, Rhodri (18 March 2016). "187-gene phylogeny of protozoan phylum Amoebozoa reveals a new class (Cutosea) of deep-branching, ultrastructurally unique, enveloped marine Lobosa and clarifies amoeba evolution". Molecular Phylogenetics and Evolution. 99: 275–296. Bibcode:2016MolPE..99..275C. doi:10.1016/J.YMPEV.2016.03.023. PMID27001604.
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