Taxonomy of Protista

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Protista taxonomy vs phylogeny.png
Comparison between the taxonomy and the phylogeny of protists. Notice the paraphyly of the kingdoms and subkingdoms

A protist (/ˈprtɪst/) is any eukaryotic organism (one with cells containing a nucleus) that is not an animal, plant, or fungus. The protists do not form a natural group, or clade, since they exclude certain eukaryotes with whom they share a common ancestor;[a] but, like algae or invertebrates, the grouping is used for convenience. In some systems of biological classification, such as the popular five-kingdom scheme proposed by Robert Whittaker in 1969, the protists make up a kingdom called Protista, composed of "organisms which are unicellular or unicellular-colonial and which form no tissues".[1][2][b]

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:

* = Paraphyletic group
† = Extinct group
? = Group no longer mentioned in recent taxonomical publications

Kingdom Protozoa*

The Protozoa Godfuss (1817) is a paraphyletic assemblage of several groups of heterotrophic microorganisms. They are divided into two main groups:[3]

Subkingdom Sarcomastigota* Cavalier-Smith (1983), which contains the protist phyla more closely related to fungi and animals: Amoebozoa, Choanozoa and Sulcozoa*, as well as the phylum Microsporidia that has recently been integrated into the Fungi kingdom instead.[4]
Subkingdom Eozoa* Cavalier-Smith (2003), which contains the excavate phyla Euglenozoa, Percolozoa, Loukozoa* and Metamonada.

Subkingdom Sarcomastigota*

Phylum Amoebozoa

Phylogeny of Amoebozoa[5]
Amoebozoa

Discosea

Tevosa

Tubulinea

Evosea

Cutosa

Conosa

Archamoebea

Semiconosia

Variosea

Mycetozoa

The current classification of phylum Amoebozoa Lühe 1913 em. Cavalier-Smith 1998 isn't entirely phylogenetic: subphylum Lobosa* is a paraphyletic group (Glycopoda* and Cutosa) as opposed to the monophyletic subphylum Conosa, and the superclass Glycopoda* is also a paraphyletic grouping (Discosea and Tubulinea) as opposed to the monophyletic superclass Cutosa. Phylogenetic trees show that Discosea is sister to Tevosa (Tubulinea + Evosea), and Tubulinea is sister to Evosea (Cutosa + Conosa).[5]

Phylum Choanozoa*

Phylogeny of Choanozoa*[4][15]
Opisthokonta
Holomycota
Fungi

Eumycota

Aphelida

Rozellida

Microsporidia

Cristidiscoidea

Holozoa

Ichthyosporea

Pluriformea

Filozoa

Filasterea

Apoikozoa

Choanoflagellatea

Metazoa

The phylum Choanozoa Cavalier-Smith 1981 is a paraphyletic grouping of all protists in the clade Opisthokonta that aren't fungi or animals. It is divided into two subphyla: Choanofila, more closely related to Kingdom Animalia, and Paramycia, more closely related to Kingdom Fungi.[16] Although previously grouped in Paramycia, recent taxonomies place Aphelida and Rozellida as separate phyla inside the Fungi kingdom.[4]

Subkingdom Eozoa*

Phylum Loukozoa

Class Jakobea Cavalier-Smith (1999)
Order Jakobida
Family Histionidae
Family Jakobidae
Class Malawimonadea
Order Malawimonadida
Family Malawimonadidae

Phylum Metamonada

Phylum Percolozoa

Phylum Euglenozoa

Kingdom Chromista

Subkingdom Hacrobia*

Phylum Telonemia

Phylum Haptista

Phylum Cryptista

Subkingdom Harosa

Rhizaria

Phylogeny of Rhizaria[35]
Rhizaria
Cercozoa
Filosa

Reticulofilosa

Monadofilosa

Endomyxa (taxonomically in Retaria)

Lapot gusevi

Retaria

Foraminifera

Polycystinea

Acantharea

The taxonomy of Rhizaria Cavalier-Smith, 2002 has changed in recent years due to the change in placement of Endomyxa. Originally, phylum Cercozoa Cavalier-Smith, 1998 was made up of two subphyla: Filosa and Endomyxa.[36] The taxonomy of Cercozoa was then revised in 2018, after phylogenetic analyses placed Endomyxa in phylum Retaria Cavalier-Smith, 2002 instead of Cercozoa, rendering the taxon Filosa as a synonym for Cercozoa.[19] However, thanks to posterior phylogenomic analyses in 2019, the monophyly of Cercozoa (Filosa + Endomyxa) was confirmed, meaning Endomyxa isn't phylogenetically part of Retaria.[35]

This article follows the latest taxonomy, where Endomyxa is still part of Retaria and not Cercozoa,[19] despite this not being phylogenetically true.[35]

Phylum Cercozoa

Phylum Retaria

Phylogeny of Retaria[38]
Retaria
Foraminifera
Globothalamea

Rotaliida

"Textulariida"

"Monothalamea"

Tubothalamea

"Monothalamea"

Polycystinea

Acantharea

Taxopodida

Superphylum Alveolata

Phylum Ciliophora

Phylum Myzozoa

Superphylum Heterokonta

Phylum Bigyra

Phylogeny of Bigyra[46][47]
Bigyra
Opalozoa
Placidozoa
Opalinata

Opalinea

Blastocystea

Opalomonadea  

Nanomonadea

Placididea

Bikosea

Sagenista

Eogyrea

Labyrinthulea

Phylum Gyrista

Phylogeny of Gyrista[47][29][52]
Gyrista
Ochrophyta
Chrysista
“SI clade”
Fucistia

Xanthophyceae

Chrysoparadoxophyceae

Phaeosacciophyceae

Schizocladiophyceae

Phaeophyceae

Aurophyceae

Aurearenophyceae

Phaeothamniophyceae

“PX clade”
Raphidoistia

Raphidophyceae

Limnistia

“SII clade”
Diatomista

Hypogyrista

Khakista

Bolidophyceae

Diatomea

“SIII clade”

?Bigyromonada

Pseudofungi

Oomycetes

Hyphochytrea

Kingdom Archaeplastida

Despite being technically inside the plant kingdom, rhodophytes and glaucophytes are considered protists under some definitions.

Division Glaucophyta

Division Rhodophyta

Protista incertae sedis

Meteora sporadica Hausmann, Weitere, Wolf & Arndt (2002)[66]

See also

Notes

  1. ^ The first eukaryotes were “neither plants, animals, nor fungi”, hence as defined, the Protista would include the earliest common ancestor of all eukaryotes.
  2. ^ In the original 4-kingdom model proposed in 1959, Protista included all unicellular microorganisms such as bacteria.
  3. ^ These last three genera are considered as "possible" Centrohelea.[28]
  4. ^ The heliomonads, including Heliomorpha, have been transferred back into Cercozoa in an unpublished paper submitted in 2020 that revises the classification of Cercozoa. It is thought that the transnuclear cytoplasmic channels for the axopodial axonemes found both in Microhelida and Heliomonadida evolved independently.[32]
  5. ^ Phylogenetically, Endomyxa belongs to Cercozoa, not Retaria,[35] but the most recent taxonomy places it in Retaria.[19]
  6. ^ Probably paraphyletic.
  7. ^ This paraphyletic class includes the orders Allogromiida and Astrorhizida, the Xenophyophorea, and several unnamed clades recovered from environmental sampling.[44]
  8. ^ The new classification[45] only includes genera for which there is molecular data; the rest of the taxa contained in the older classification[43] follow a question mark "?".
  9. ^ This paraphyletic grade includes the polyphyletic Lituolida, the Loftusiida and the Textulariida sensu stricto.[44]
  10. ^ Polyphyletic group.[44]
  11. ^ This genus is currently the earliest diverging lineage of Labyrinthulea, which is why it still doesn't belong to any family.[49]
  12. ^ Though previously part of the order Thraustochytrida,[50] it is phylogenetically outside of any known order of Labyrinthulea and is the second earliest diverging lineage after Stellarchytrium.[49]
  13. ^ Probably paraphyletic.[29]
  14. ^ Cavalier-Smith suggested that Sulcochrysis is possibly related to Olisthodiscus, and grouped both genera within this subclass.[46][29] However, a study in, 2021 reveals that Olisthodiscales is part of a new class Olisthodiscophyceae, which includes only the genus Olisthodiscus. The position of the genus Sulcochrysis remains uncertain due to a lack of DNA sequences.[65]

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