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This page is about the present classification of protists. For the science of naming and classifying protists, and its history, see Protist systematics.

This is a test page for improving the Protist classification article.

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Legend:

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

Overview of classification

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Cladogram of eukaryotes showing the major protist groups, based on 2023–2025 phylogenetic analyses.

Eukarya

Amorphea

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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.[1]

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.[2] 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.[3][4]

Amoebozoa

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Cladogram of the major groups of Amoebozoa based on an 824-gene phylogenetic analysis published in 2022.[5]

The phylum Amoebozoa contains around 2,400 species[6] of primarily amoeboid protists.[7] 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.[8] After the general 2019 revisions published by the International Society of Protistologists (ISOP),[7] there have been specific revisions to the classification of eumycetozoans[9] and testate amoebae.[10]

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.[7]

Obazoa

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Cladogram of Obazoa based on several analyses.[29][30][31]

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.[7] Under the Cavalier-Smith system, breviates and apusomonads were two classes that composed the phylum Apusozoa,[32] but this taxon is paraphyletic, as apusomonads are more closely related to opisthokonts.[29] The taxonomy of apusomonads was expanded in a 2022 phylogenetic sudy that introduced many new genera.[33]

Obazoa Brown et al. 2013

Opisthokonta

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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[7] and mycologists alike.[27] Both Holozoa and Nucletmycea have been proposed once as superkingdoms by a group of mycologists, specifying nucleariids and fungi as kingdoms, but without any mention of holozoan kingdoms.[34]

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

Archaeplastida

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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).[32] As such, the protist lineages, including simple, single-celled algae and protozoa, are regarded as "plants" under this definition.[44] Other definitions exclude these protists, and reduce the plant kingdom to only embryophytes.[7]

Rhodophyta

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Cladogram showing the relationships between red algal classes, according to a 2017 phylogenomic analysis.[45]

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.[46] Unlike other eukaryotes, they lack flagella and centrioles in all their life stages.[47]

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.[48] 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.[49] This taxonomic system, developed by Hwan Su Yoon and collaborators, is the prevalent among taxonomists,[50][46] with the addition of a separation of subphylum Rhodophytina into two subphyla Proteorhodophytina and Eurhodophytina in 2017.[45] In 2022, a book was published revising the classification of all freshwater red algae.[51]

Phylum Rhodophyta Wettstein 1901

Rhodelphidia

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Picozoa

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Glaucophyta

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Chloroplastida

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Cladogram of Chloroplastida with focus on the classes of green algae, based on 2019–2023 phylogenetic studies. Prasinophytes are marked *.[77][78][79]

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

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Division Prasinodermophyta Marin & Melkonian in Li et al. 2020[78]

Chlorophyta

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

Streptophyta

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

Pancryptista

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Cladogram of Pancryptista based on 2020 and 2022 analyses.[92][93]

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,[94] the algal division described by the same author in previous years.[95] 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.[96] The next year, a separate group of authors proposed a different higher ranking for katablepharids as a phylum Kathablepharida or division Katablepharidophyta, related to but independent from phylum Cryptophyta, with no mention of Cryptista.[97] 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;[98] and Corbihelia, containing picozoans, telonemids, and some heliozoan species (Heliomorpha and Microheliella).[99] Corbihelia did not reach consensus: later analyses and revisions separated telonemids and picozoans as their own clades, and placed Heliomorpha in Cercozoa.[7] However, the addition of Palpitomonas and the monophyly of Rollomonadia have been supported in other analyses.[92]

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,[7] 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[100] and often excluded goniomonads too.[99] 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.[93]

Pancryptista Yazaki et al. 2022

Haptista

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Alveolata

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Ciliophora

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Myzozoa

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Chrompodellids

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Apicomplexa

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Perkinsozoa

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Dinoflagellata

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Rhizaria

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Cercozoa

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Endomyxa

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Foraminifera

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Globothalamea

Xenophyophorea

'monothalamids'

'monothalamids'

Tubothalamea

'monothalamids'

Cladogram of foraminifera based on a 2022 phylogenetic analysis.[104]

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.[105] 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,[106] until they were placed as a subphylum of phylum Retaria along with the Radiolaria.[101] They are divided into three classes (or subclasses, when globally treated as a class): the paraphyletic monothalamids,[107] 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.[104] Some alternate systems of dividing foraminifera into classes have been proposed,[108] but none have gained consensus. The relationships of monothalamids remain a work in progress due to the limited amount of genetic data.[104]

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.[109] 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[106] and 2014,[110] and one classification for planktonic foraminifera was published in 2022.[111]

Subphylum Foraminifera d'Orbigny 1826

Radiolaria

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Subphylum Radiolaria Müller 1858 sensu Adl et al. 2005

Stramenopiles

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Cladogram of stramenopiles, based on 2022–2024 phylogenomic analyses.[126][127]

The stramenopiles are a diverse group of protists composed of more than 100,000 estimated species.[128] They are generally biflagellates, characterized by straw-like tubular flagellar hairs in one of their flagellum; this is referenced in their name (Latin stramen, 'made of straw').[129] They are also known as heterokonts due to their unequal flagella, as only one of them has hairs.[130] The formal taxonomic name Heterokonta was originally proposed by Cavalier-Smith as a small phylum of algae (namely the chrysophytes, xanthophytes, phaeophytes and raphidophytes).[131] Its definition became progressively wider to include diatoms, eustigmatophytes, and non-algal groups like the oomycetes,[132] and it became a superphylum within his kingdom Chromista.[101] Simultaneously, the name 'stramenopile' was coined by David Patterson for the same group of organisms[133] and reached wide usage in the scientific community, as it was more descriptive of their characteristic flagella,[129][7] while the term 'heterokont' could also refer to any protist with flagella of unequal size (e.g., euglenids, jakobids).[134] In the 2005 ISOP revision of protist classification, the term was formalized as a taxon with a phylogenetic definition and was capitalized as 'Stramenopiles'.[129] Variants of this name have been proposed in different ranked systems, such as kingdom 'Straminipila'.[135][136]

Most stramenopiles are classified into two phyla: Bigyra, containing the earliest-branching heterotrophs, and Gyrista, containing the photosynthetic stramenopiles (Ochrophyta) and some of their heterotrophic relatives, like oomycetes. Bigyra is further divided into two subphyla: Sagenista, containing the labyrinthulids, and Opalozoa, containing the bicosoecids, opalinids. In addition, the earliest branching stramenopile, Platysulcus tardus, has been proposed as incertae sedis Bigyra.[101] However, most taxonomists exclude Platysulcus from this phylum.[7][137] Another basal stramenopile, Kaonashia, was described without explicit assignment to any taxon within stramenopiles.[138]

Stramenopiles Patterson 1989 emend. Adl et al. 2005 (=Heterokonta Cavalier-Smith 1986; Straminipila Dick 2001)

Bigyra

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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.[139][101]

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.[7][126][127] Its global classification was last revised by Cavalier-Smith in 2013 and 2018,[98][101] while the specific class Placididea was revised in 2021, with new taxa described by Alexandra Rybarski and coauthors,[140] and the classes Labyrinthulea and Opalinea were revised in-depth in the 2017 Handbook of the protists.[141][142]

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

Bigyromonada

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Subphylum Bigyromonada Cavalier-Smith 1998

Pseudofungi

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The pseudofungi are a group of over 1,200 species[148] of fungus-like protists, composed of two groups: the hyphochytrids and oomycetes. Both groups have been traditionally studied by mycologists and regarded as phylum-level taxa, Hyphochytriomycota and Oomycota (or Peronosporomycota), inside the kingdom Straminipila.[135][136] Some protistologists place them as classes of the phylum or subphylum Pseudofungi instead.[101] Due to their similarity to fungi, they (and labyrinthuleans) are also known as 'straminipilous fungi'.[135] Their classifications were reviewed in the 2017 work Handbook of the protists. The taxonomy of oomycetes is provisional, as molecular phylogenetics are still resolving the inner relationships.[149][7]

Subphylum Pseudofungi Cavalier-Smith 1986 (=Heterokontimycotina M.W. Dick 1976, Peronosporomycotina M.W.Dick 2001;[135]: 288  Pythiista Cavalier-Smith 1986)[132]

Ochrophyta

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Evolutionary relationships between all ochrophyte classes, based on a 2024 multiprotein phylogenetic analysis.[151]

The ochrophytes (also known as heterokontophytes or heterokont algae) are a vast group of mostly photosynthetic stramenopiles.[152] The first phylum name introduced specifically for heterokont algae was Heterokontophyta, coined by van den Hoek in 1978 and subsequently used by several authors,[153] but it was not validly published until 2023.[154] Meanwhile, T. Cavalier-Smith coined in 1986 the alternative name Ochrista, renamed Ochrophyta in 1996 to match the recommendations of the International Code of Botanical Nomenclature.[153] The same author later renamed it Ochrophytina after making it a subphylum of phylum Gyrista,[101] but the scientific community continues to study heterokont algae as a phylum Ochrophyta or Heterokontophyta.[7][152]

The numerous classes of ochrophytes are currently organized into two major clades: Chrysista, containing the clades SI and SII, and Diatomista, equivalent to the clade SIII. However, the exact relationships of ochrophyte classes within these smaller clades continue to be debated.[152]

Subphylum/Phylum Ochrophyta Cavalier-Smith 1986 emend. Cavalier-Smith & Chao 1996 (=Heterokontophyta Guiry, R.A.Andersen & Moestrup 2023). Genera incertae sedis: Commation,[m] Chrysoderma*, Chrysomeris*, Chrysonephos*, "Giraudyopsis",[157] Rhamnochrysis*.[n]

Diatoms

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Further information: Taxonomy of diatoms

The diatoms (clade Diatomeae)[7] are a particularly species-rich group of stramenopiles, with more than 14,000 described species.[196] They are often classified as an entire phylum or division Bacillariophyta by diatomists, despite being part of the phylum Ochrophyta.[197] Taxonomists divide diatoms into three main classes: Coscinodiscophyceae, for radial centric diatoms; Mediophyceae, for polar centric diatoms; and Bacillariophyceae, for pennate diatoms.[198] However, the current class- and order-level classifications do not reflect molecular phylogenetics: most classes and many orders are non-monophyletic, and some orders are spread between different classes.[199][200] The ISOP publication of 2019 proposed a newer preliminary classification with higher ranks to reflect the advances in phylogenetics of the 2010s decade, although it still suffers from poor taxon sampling outside of the polar centric and pennate diatoms, and expects the basal taxa to become better resolved in the future.[7]

Diatomeae Dumortier 1821 (=Bacillariae Haeckel 1878, Bacillariophyta Karsten 1928)

Excavates

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Malawimonada

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Metamonada

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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).[203] These were later split into other groups such as the Parabasalia and Preaxostyla (over 260 and 140 species respectively),[204][205] 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;[206][207] around 140 species).[204] In this system, Preaxostyla is known as subphylum Anaeromonada, and Fornicata and Parabasalia are two infraphyla that belong to the subphylum Trichozoa.[32] Still, some taxonomists retain the Parabasalia at a phylum level.[208]

The classifications of Preaxostyla and Fornicata were last revised separately in the 2017 Handbook of the protists[209][210][211] and the higher classification of Metamonada was revised by Cavalier-Smith in 2021.[32] The classification of Parabasalia was completely updated in 2024.[208] Two new smaller clades of metamonads have been described in addition: anaeramoebae[212] and the 'BaSk' clade, containing barthelonids and skoliomonads.[213] However, Parabasalia was treated as a separate phylum by the 2024 revision, instead of a member of phylum Metamonada.[208] For consistency, the classification listed below maintains Parabasalia as part of the phylum Metamonada.

Phylum Metamonada Grassé 1952 emend. Cavalier-Smith 2003

Discoba

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Jakobida

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Heterolobosea

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Cladogram of heterolobosean orders based on combined phylogenomic and 18S rRNA analyses published in 2025.[218]

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,[7] 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.[219][220] The first comprehensive phylogenomic study of Heterolobosea was published in 2025 by Tomáš Pánek and coauthors, resulting in its modern classification.[218]

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

Euglenozoa

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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.[221][222][223] 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.[223] The classification of euglenozoans was summarized by Cavalier-Smith in 2016.[222] A more phylogenetically precise revision of their classification was published in 2021 by Alexei Kostygov and coauthors,[223] with newer clades and genera described in the following years, particularly of euglenids. Still, one group of euglenids, the paraphyletic "ploeotids",[224] remains unresolved in the current classification, spread out across multiple clades.[225][226][227]

Phylum Euglenozoa Cavalier-Smith 1981 emend. Simpson 1997

Minor clades

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Ancyromonadida

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CRuMs

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Hemimastigophora

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Provora

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Telonemia

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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.[236] Under the Cavalier-Smith system, telonemids were initially classified as a class of Cryptista,[99] but later analyses consistently recovered it as a separate group.[32] Until 2019, only two species had been formally described, each belonging to a separate genus,[237] although environmental DNA sequencing suggests there are many more species not yet described.[238] In 2022, five additional species were described along with a third new genus, bringing the total number of species to seven.[239]

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

Protists of uncertain position

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Notes

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  1. ^ 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.[7]
  2. ^ This genus grouped with Dermamoebidae in a phylogenetic anaysis that used a limited number of taxa.[18] In the recent taxonomy it was listed as a separate clade until its position was better resolved.[7] A 2023 analysis placed it, once again, related to Dermamoebida in a clade with Microglomus.[19]
  3. ^ The two genera of Stygamoebidae, Stygamoeba and Vermistella, group together in some phylogenetic analyses, but usually appear separated. In 2019 they were considered only a potential branch of Discosea, unless proven otherwise.[7] A 2023 analysis places both genera in Flabellinia, closely related to Thecamoebida.[19]
  4. ^ a b c d e The orders Thecamoebida, Dactylopodida, Acanthopodida, Himatismenida and Pellitida were initially divided into families, but phylogenetic analyses haven't supported the monophyly of each family. For this reason, in recent classifications there is no family rank division within it.[11][7]
  5. ^ The genus Pessonella could be a synonym of Vannella.[7]
  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.[7]
  7. ^ a b According 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."[68]
  8. ^ The 2019 revision by the ISOP wrongly stated that goniomonads are classified as "Cyathomonadacea Pringsheim 1944".[7] 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.[100]
  9. ^ The genus Tetragonidium is known only from one written diagnosis and some illustrations, and its affinities with cryptomonads are very uncertain.[100]
  10. ^ 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.[123]
  11. ^ 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.[98] However, both of those families remain accepted by the scientific community as independent from Cafeteriaceae as of 2020.[143]
  12. ^ In the Handbook of the protists, taxonomists Gordon Beakes and Marco Thines assume oomycetes are a phylum-level taxon (Oomycota), and raise the subclasses Saprolegniomycetidae and Peronosporomycetidae to classes Saprolegniomycetes and Peronosporomycetes, respectively.[149] However, most taxonomists treat the oomycetes as a class-level taxon, as represented here.[101][7]
  13. ^ a b Cavalier-Smith hypothesized in 2013 a sister relationship between actinophryids and raphidophytes, with both groups placed under the taxon Raphidomonadea. The molecularly uncharacterized genus Commation was classified as part of this taxon. [98] However, the placement of actinophryids among ochrophytes is still unstable and uncertain,[155][156] and Commation has never been included in classifications since.[7]
  14. ^ The eight genera Antarctosaccion, Chrysoderma, Chrysomeris, Chrysonephos, Chrysowaernella, Giraudyopsis, Phaeosaccion and Rhamnochrysis were initially considered golden algae (class Chrysophyceae). In 1989, Charles O'Kelly claimed they had been classified in a new order Chrysomeridales according to an unpublished article.[158][159] Later, Thomas Cavalier-Smith placed this unpublished order inside the class Chrysomeridophyceae.[160] However, the class was proven to be polyphyletic: Chrysowaerella was transferred to Chrysoparadoxophyceae, while Antarctosaccion and Phaeosaccion were moved to Phaeosacciophyceae. Due to unavailable molecular data, the placement of Chrysoderma, Chrysomeris, Chrysonephos and Rhamnochrysis remains unknown,[161] although Chrysonephos possibly belongs to the Sarcinochrysidales.[162][163]
  15. ^ The position of Eustigmatophyceae is still unstable, with different phylogenetic methodologies showing affinities to either of the SI and SII clades.[165][156]
  16. ^ In Syllabus of plant families (2015),[174]: 126  the chrysophyte family Chrysosphaeraceae (containing the genera Chrysosphaera, Chrysastrella and Epicystis) is considered incertae sedis with no assigned order. Later publications place this family (or at least its genera) within the order Chrysosaccales,[175][176] but maintain one of its genera, Chrysastrella, as incertae sedis among chrysophytes.[170]
  17. ^ According to the Syllabus of plant families (2015), the order Ochromonadales contains three families, Chrysolepidomonadaceae, Dinobryaceae, and Ochromonadaceae. However, these are not monophyletic,[174]: 122–123  and many phylogenetic studies of the following years have abandoned the use of families.[7][178][169][179]
  18. ^ Some authors consider Ciliophryales and Rhizochromulinales to be separate orders.[192]
  19. ^ Although formally recognized as a genus of Sarcinochrysidales, Ankylochrysis groups with Pelagomonadales in phylogenetic analysis.[193]
  20. ^ One 18S rRNA phylogenetic analysis suggests that the genus Opisthomitus is affiliated with the family Pyrsonymphidae.[209]
  21. ^ Regarding the classification of Fornicata, there is a disconnect between the Cavalier-Smith system and the system described by different authors in the Handbook of the protists and other studies. The order Retortamonadida was originally defined as two genera: Chilomastix and Retortamonas. As both groups of taxonomists have reported, retortamonads are polyphyletic, because Chilomastix branches distantly from Retortamonas. In 2013, Cavalier-Smith fixed this taxonomic issue by modifying the retortamonads to only include Retortamonas, and describing a separate order Chilomastigida for Chilomastix only, placed among the early branching Carpediemonas-like organisms.[214] However, this has not yet been recognized by later studies from the other authors, as they have continued to report a polyphyletic Retortamonadida.[211][215][216] Because of this inconsistency, Eopharyngea (the clade formed by Retortamonadida plus Diplomonadida) is monophyletic only when considering one version of Retortamonadida, but not the other.
  22. ^ The family Orodruinidae is possibly polyphyletic, and its type genus Orodruina may belong to the order Neovahlkampfiida.[218]

References

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