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Protist classification

Classification of eukaryotes

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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; but, like algae or invertebrates, the grouping is used for convenience. In some systems of biological classification, such as the 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". In the 21st century, the classification shifted toward a phylogenetic classification of protists.

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

01Cavalier-Smith's classification

Thomas Cavalier-Smith and his collaborators use a fully rank-based classification of protists. Here is their classification:

Extended content

02Current classification

Legend:

Extinct, or exclusively fossil taxon.
(P) Paraphyletic or polyphyletic taxon.
(P?) Potentially paraphyletic or polyphyletic taxon.
(=...) Taxonomic synonym.
(...) Same taxon in a different code of nomenclature.

03Amorphea

The supergroup Amorphea contains very diverse heterotrophic organisms, from the macroscopic fungi and animals to the unicellular choanoflagellates and classical amoebae. They frequently exhibit the ability to produce multinucleated cells, a trait considered ancestral to Amorphea. They are also capable of producing pseudopodia, as does the closely related CRuMs clade, forming the clade Podiata.

Amorphea is divided into two clades: Amoebozoa, containing well-known amoebae and slime molds, and Obazoa, containing animals, fungi, and their closest relatives. The relationship between these two clades was initially called 'Unikonta', due to a hypothesis where their common ancestor was a unikont, i.e., a eukaryote with just one flagellum. However, this hypothesis was refuted, as there are bikont amorpheans (e.g., Breviata anathema) and it likely is not an ancestral trait to have a single flagellum.

Obazoa

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

Opisthokonta

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

Amoebozoa

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

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

04Archaeplastida

The clade Archaeplastida encompasses photosynthetic eukaryotes whose chloroplasts are derived from an event of primary endosymbiosis with a cyanobacterium. This includes the multicellular land plants (embryophytes), distinguished by their embryonic development and differentiation into tissues, and a multitude of protist lineages, many of which have also evolved multicellularity independently (e.g., red algae). Together, they amount to an estimated 500,000 species, with protists covering less than 50,000. In some systems of classification, Archaeplastida is equivalent to the plant kingdom (Plantae). As such, the protist lineages, including simple, single-celled algae and protozoa, are regarded as "plants" under this definition. Other definitions exclude these protists, and reduce the plant kingdom to only embryophytes.

Rhodophyta

Cladogram showing the relationships between red algal classes, according to a 2017 phylogenomic analysis.

The red algae, formally known as phylum Rhodophyta, are a group of more than 7,100 species of photosynthetic eukaryotes containing primary plastids. They have evolved into diverse morphologies, from single coccoid cells to large pseudoparenchymatous thalli that have complex life cycles with up to three generations. Unlike other eukaryotes, they lack flagella and centrioles in all their life stages.

Traditionally, red algae have been classified according to morphology as two classes (Bangiophyceae and Florideophyceae), or two subclasses (Bangiophycidae and Florideophycidae) of a single class Rhodophyceae. This system was challenged with the rise of molecular phylogenetics, which revealed the paraphyly of Bangiophyceae. In addition, a third class, Cyanidiophyceae, was discovered as the earliest diverging lineage. In 2004 a new molecular-based classification of red algae was proposed, where they composed a subkingdom of plants (Rhodoplantae) divided into two phyla: Cyanidiophyta, with just one class, and Rhodophyta, with three subphyla (Rhodellophytina, Metarhodophytina and Eurhodophytina) and four classes, one of which was still paraphyletic. In 2006, a comprehensive molecular analysis revised the classification and proposed a single phylum Rhodophyta with two subphyla: Cyanidiophytina, with one class, and Rhodophytina, with six classes. This taxonomic system, developed by Hwan Su Yoon and collaborators, is the prevalent among taxonomists, with the addition of a separation of subphylum Rhodophytina into two subphyla Proteorhodophytina and Eurhodophytina in 2017. In 2022, a book was published revising the classification of all freshwater red algae.

Phylum Rhodophyta Wettstein 1901

Rhodelphidia

Phylum Rhodelphidia Tikhonenkov, Gawryluk, Mylnikov & Keeling 2019 → class Rhodelphidea Tikhonenkov, Gawryluk, Mylnikov & Keeling 2019 → order Rhodelphida Tikhonenkov, Gawryluk, Mylnikov & Keeling 2019 → family Rhodelphidae Tikhonenkov, Gawryluk, Mylnikov & Keeling 2019. Sole genus: Rhodelphis Tikhonenkov, Gawryluk, Mylnikov & Keeling 2019.

Picozoa

Phylum Picozoa Seenivasan, Sausen, Medlin & Melkonian 2013 (=picobiliphytes Not et al. 2007) → class Picomonadea Seenivasan, Sausen, Medlin & Melkonian 2013 → order Picomonadida Seenivasan, Sausen, Medlin & Melkonian 2013 → family Picomonadidae Seenivasan, Sausen, Medlin & Melkonian 2013. Sole genus: Picomonas Seenivasan, Sausen, Medlin & Melkonian 2013.

Glaucophyta

Phylum Glaucophyta Skuja 1954 (=Glaucocystaceae West 1904, Glaucocystophyta Kies and Kremer 1986, Glaucoplantae Marin & Melkonian in Li et al. 2020). Incertae sedis genera: Archaeopsis, Chalarodora, Glaucocystopsis, Peliaina, Strobilomonas.

Chloroplastida

The green algae are a paraphyletic group from which land plants (Embryophyta) evolved. Together they form the clade Chloroplastida (in reference to their green plastids), also known as Viridiplantae (meaning "green plants"). Green algae were originally studied as a single division Chlorophyta.

Prasinodermophyta

Division Prasinodermatophyta Marin & Melkonian in Li et al. 2020 (as Prasinodermophyta)

Chlorophyta

Division Chlorophyta Reichenbach 1828 emend. Pascher 1914 emend. Lewis & McCourt 2004

Streptophyta

Division Streptophyta Bremer & Wanntorp 1981 (=Charophyta Migula 1897 emend. Karol et al. 2009)

05Pancryptista

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

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

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

Pancryptista Yazaki et al. 2022

06Haptista

07Stramenopiles

  • Class Platysulcea Cavalier-Smith 2018 → order Platysulcida Cavalier-Smith 2018 → family Platysulcidae Shiratori, Nakayama & Ishida 2015. Sole genus: Platysulcus Shiratori, Nakayama & Ishida 2015.

Bigyra

Bigyra was initially proposed as a phylum of heterotrophic protists closely related to the photosynthetic heterokonts (Ochrophyta). It contained three subphyla: Pseudofungi (oomycetes and hyphochytrids), Bigyromonada (free-living biflagellates), and Opalinata (large multi-flagellated gut symbionts). The phylum was proposed with a unifying characteristic, the presence of a flagellar transition zone with double helices or rings. This definition was later modified to exclude both Pseudofungi and Bigyromonada, and it was instead reduced to two different subphyla: Opalozoa (including Opalinata and bicosoecids) and Sagenista. Prior to this, Sagenista itself was also a phylum containing bicosoecids and labyrinthuleans, but it got reduced to only labyrinthuleans and a few flagellates.

The current definition of Bigyra is paraphyletic according to some phylogenetic analyses, with Sagenista more closely related to Gyrista. Despite its probable paraphyly, most taxonomists continue to use it, and it was recognized in the 2019 revision by the ISOP, which prioritizes monophyletic taxa. Its global classification was last revised by Cavalier-Smith in 2013 and 2018, while the specific class Placididea was revised in 2021, with new taxa described by Alexandra Rybarski and coauthors, and the classes Labyrinthulea and Opalinea were revised in-depth in the 2017 Handbook of the protists.

Phylum Bigyra(P?) Cavalier-Smith 1998 emend. 2013

Gyrista

08Alveolata

Cladogram of Alveolata, based on a SSU rDNA phylogeny of basal alveolates and a 203-gene phylogeny of apicomplexans and their relatives published in 2023.

Genera not assigned to any lower taxon: Neocolponema Gigeroff et al. 2023, Loeffela Gigeroff et al. 2023.

Myzozoa

ACS

Dinozoa

Ciliophora

09Rhizaria

Cladogram of Rhizaria based on a 2019 phylogenomic analysis.

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, Rhizaria is home to over 11,000 described species.

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. Cercozoa contains two subphyla: Filosa, represented primarily by free-living lineages, and Endomyxa, which includes obligate intracellular parasites and free-living predators. Included in or related to Endomyxa are several clades known primarily from environmental sequences ('novel clades' 10, 11, 12), with only a few members formally described.

The classification below is based largely on the 2019 revision by Adl et al. and Cavalier-Smith et al. Adl et al. excludes Endomyxa and Skiomonadea from Cercozoa, but later works generally include them.

Phylum Cercozoa Cavalier-Smith 1998, emend. Adl et al. 2005. Incertae sedis genera: Bereczkya, Dictiomyxa, Myxodictyium, Pontomyxa, Protomyxa, Protogenes, Rhizoplasma.

Foraminifera

Globothalamea

Xenophyophorea

'monothalamids'

'monothalamids'

Tubothalamea

'monothalamids'

Cladogram of foraminifera based on a 2022 phylogenetic analysis.

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. 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, until they were placed as a subphylum of phylum Retaria along with the Radiolaria. They are divided into three classes (or subclasses, when globally treated as a class): the paraphyletic monothalamids, 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. Some alternate systems of dividing foraminifera into classes have been proposed, but none have gained consensus. The relationships of monothalamids remain a work in progress due to the limited amount of genetic data.

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. 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 and 2014, and one classification for planktonic foraminifera was published in 2022.

Subphylum Foraminifera d'Orbigny 1826

Radiolaria

10Disparia

Cladogram of Disparia based on a 2025 phylogenomic analysis.

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. 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. However, in 2024 there was already evidence of a close affinity between Meteora, hemimastigotes, and provorans. 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. Hemimastigotes are a small group composed of ten species of heterotrophs with a pellicle and numerous flagella. First identified in 1988, they remained an orphan group for decades due to the absence of genetic information, with taxonomists hypothesizing affinities to many different protists. In 2018, two hemimastigotes were successfully sequenced and a phylogenomic analysis revealed their deep-branching placement near Diaphoretickes. Between 2024 and 2025, a closer relationship with the clades Provora and Caelestes was resolved. 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.

11Excavates

Malawimonadida

Metamonada

The metamonads were first described by Pierre-Paul Grassé in the first volume of Traité de Zoologie, published in 1952, as the superorder Metamonadina or Anaxostylaria. They were originally composed of zooflagellates with four or more flagella, known as polymastigotes and hypermastigotes (e.g., Trichomonas, Oxymonas). These were later split into other groups such as the Parabasalia and Preaxostyla (over 260 and 140 species respectively), 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; around 140 species). In this system, Preaxostyla is known as subphylum Anaeromonada, and Fornicata and Parabasalia are two infraphyla that belong to the subphylum Trichozoa. Still, some taxonomists retain the Parabasalia at a phylum level.

The classifications of Preaxostyla and Fornicata were last revised separately in the 2017 Handbook of the protists and the higher classification of Metamonada was revised by Cavalier-Smith in 2021. The classification of Parabasalia was completely updated in 2024. Two new smaller clades of metamonads have been described in addition: anaeramoebae and the 'BaSk' clade, containing barthelonids and skoliomonads. However, Parabasalia was treated as a separate phylum by the 2024 revision, instead of a member of phylum Metamonada.

Phylum Metamonada Grassé 1952 emend. stat. n. Cavalier-Smith 2003

Discoba

Jakobida

Tsukubamonadida

Heterolobosea

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

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, 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. The first comprehensive phylogenomic study of Heterolobosea was published in 2025 by Tomáš Pánek and coauthors, resulting in its modern classification.

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

Euglenozoa

The phylum Euglenozoa is home to at least 2,000 described species of single-celled flagellates of very dissimilar lifestyles. It was originally proposed to group the euglenids (such as the photosynthetic Euglena) and the kinetoplastids (like the pathogenic Trypanosoma), usually studied separatedly; eventually it included diplonemids and symbiontids as well. 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. The classification of euglenozoans was summarized by Cavalier-Smith in 2016. A more phylogenetically precise revision of their classification was published in 2021 by Alexei Kostygov and coauthors, with newer clades and genera described in the following years, particularly of euglenids. Still, one group of euglenids, the paraphyletic "ploeotids", remains unresolved in the current classification, spread out across multiple clades.

Phylum Euglenozoa (Euglenophyta) Cavalier-Smith 1981 emend. Simpson 1997

12Minor clades

Ancyromonadida

CRuMs

Telonemia

The phylum Telonemia (telonemids) contains a few species of flagellates found in ocean and fresh waters worldwide. It was originally proposed in 2006 for Telonema, a genus of previously uncertain affinity. Under the Cavalier-Smith system, telonemids were initially classified as a class of Cryptista, but later analyses consistently recovered it as a separate group. Until 2019, only two species had been formally described, each belonging to a separate genus, but environmental DNA sequencing suggested there were many more species not yet described. Between 2022 and 2025, eight additional species were described along with three new genera, bringing the total number of species to ten.

13Protists of uncertain affiliation

Incertae sedis within Diaphoretickes:

Incertae sedis within SAR:

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Sources and credits

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