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This glossary provides an overview of terms used in the description of protists, eukaryotic organisms that are neither animals nor plants nor fungi, as well as their life cycles, feeding mechanisms, and relationship with the environment. The term "protist" embraces all organisms that descended from the last eukaryotic common ancestor except those three "higher" kingdoms of life; as such, protists usually follow the same basic principles of biology as them.[1] Nevertheless, protists exhibit almost all of the spectrum of biological characteristics expressed in eukaryotes, including many unique adaptations that are covered here.[2][3]
Organism that thrives in oxygen-rich environments. Contrast: anaerobe.[4]
agamont
Life phase formed through the development of the zygote, which in turn undergoes meiosis to produce gametes. Contrast: gamont.[5]
aggregative multicellularity
Also aggregative fruiting and aggregation. Behavior of certain slime molds (e.g., dictyostelids, acrasids) consisting of many individual cells aggregating together to form the fruiting bodies, known as sorocarps.[6][7] It is considered a separate type of multicellularity from clonal multicellularity.[8]
Also amoebomastigote. Term usually reserved for single-celled protists that, during their life cycle, develop a flagellate life stage and a separate amoeba stage (as in Naegleria), with the latter lacking basal bodies.[12] Protists that have both flagella and pseudopodia in the same life stage (as in Cercomonas) are usually referred to as amoeboid flagellates instead.[13] However, the terms are sometimes used interchangeably.[14]
Any member of Amoebozoa, a phylum of around 2,400 species[15] containing many of the classical amoebae and many slime molds.[6] They are ancestrally biflagellates, but many lost one or both flagella.[16] The group is closely related to obazoans, together forming the clade Amorphea.[17]
Any member of Amorphea, the clade that groups obazoans and amoebozoans.[17] Among amorpheans, the ability to produce multinucleated cells is particularly frequent, and is considered their ancestral trait. The CRuMs clade is closely related to amorpheans; the two groups compose the clade Podiata.[16]
Organism that occupies low-oxygen environments, such as the animal gut and aquatic sediments. Anaerobic protists (e.g., metamonads) tend to evolve a drastically different mitochondrial metabolism. Contrast: aerobe.[4]
Any member of Apicomplexa, a group of more than 6,000 species of single-celled, parasitic protists that use an apical complex to penetrate cells of their animal hosts. Their life cycle consists of a double or triple-phase alternation of generations, each with typical infection, growth, and multiplication stages. In coccidians and haemosporidians, the stages are: sporogony, where a zygote differentiates into sporocysts that produce sporozoites, which penetrate cells; merogony, where the sporozoites differentiate into meronts and produce merozoites; and gamogony, where some merozoites differentiate into gamonts and begin producing gametes. In other apicomplexans there is no merogony, and sporozoites differentiate into meronts directly.[22][23]
Highly reduced plastid found in apicomplexans;[23] its function may be related to the metabolism of fatty acids.[22] It is the most intensely studied organelle of sporozoites.[23]
Also Archaezoa. Refers to amitochondriate protists; used in older hypotheses where these protists were considered more primitive than those with mitochondria. Such hypotheses were later disproved, as these protists evolved from mitochondrion-bearing ancestors.[10] Meaning "original creatures", the term was initially a synonym for protozoa, coined in 1852 by Maximilian Perty.[22]
Specialized cell in the life cycle of diatoms that is formed by differentiation of the zygote: it produces an organic wall and expands to restore the maximum size characteristic of the diatom species, often inserting silica elements (namely the incunabula and the perizonium) into the wall during this process.[26]
Also bradyzoic merozoite. A merozoite characterized by sessile, slow growth and replication, present in a chronic coccidial infection. Contrast: tachyzoite.[28]
Plural cilia. Synonym of eukaryotic flagellum or undulipodium. Its usage highly depends on the author: some reserve it for shorter appendages, and use flagellum for longer ones,[32][33][34] while others use it for all eukaryotic flagella.[35]
clonal multicellularity
Development of most multicellular organisms (e.g., animals, plants, fungi) where their multi-celled forms arise from a series of dividing cells that remain closely connected and, through additional cell division, may differentiate into further cell types. It contrasts with aggregative multicellularity, where multi-celled structures arise from the clustering of cells that otherwise reproduce separately. Among protists, clonal multicellularity is displayed in a variety of distantly related clades (e.g., green algae, red algae, brown algae, golden algae, oomycetes, ichthyosporeans). In a few cases it is facultative, as in the colonies of choanoflagellates.[36]
Plural cristae. Folds found in the inner membrane of mitochondria. The shape of these folds has been used as a taxonomic character to distinguish between major groups of eukaryotes (see discicristate).[39]
Cryptomonad cultureAlso cryptophyte. Any member of Cryptomonada, a group of aquatic flagellatedalgae comprising more than 100 species. In botanical nomenclature, they are known as the algal division Cryptophyta.[42] Their chloroplasts were obtained through endosymbiosis with a red alga.[43] Their ejectisomes are composed of two coiled ribbons or "scrolls", as opposed to their closest relatives katablepharids which have ejectisomes of one scroll each.[17]
Non-motile, resistant life stage that develops in response to stressful environmental conditions as a survival mechanism, allowing the organism to persist (resting cyst) or leading to the production of gametes (sexual cyst).[5]
Refers to a unique generation of pseudopodia consisting of sudden outwards and lateral bulging during locomotion; characteristic of heterolobosean amoebae.[12]
Plural flagella. Motile appendage present in flagellates and some prokaryotes. In the context of eukaryotes, it is synonymous with undulipodium, which excludes prokaryotic flagella.[32][33][34]
fruiting
The process of forming a fruiting body, a "fungus-like" reproductive structure that raises spores (or a sporangium) above the substrate for dispersal during the life cycle of slime molds. They can either be sorocarps, when formed through the aggregation of many cells,[48] or sporocarps, when formed through the growth of a single cell.[6]
Plural gametangia. Specialized cell or multicellular structure that produces gametes for sexual reproduction. If reproduction is oogamous, the gametangia are differentiated into female (oogonia) and male (antheridia or spermatogonia).[49][5]
In flagellates, movement that is closely associated to the surface, as opposed to swimming; flagellates usually glide on either the flagella or the cell body.[31]
Any member of Haemosporidia, one of the two groups of hematozoans within the apicomplexans. They compose around 500 species, including the malarial parasites.[23]
Any member of Holomycota or Nucletmycea, the opisthokont clade that contains fungi and their closest protist relatives, nucleariids; excludes animals and their closest relatives, collectively known as holozoans. The clade was first described as Nucletmycea, but Holomycota became a more popular name.[39]
Any member of Holozoa, one of the two opisthokont clades, containing animals and their closest protist relatives; excludes fungi and their closest relatives, collectively known as holomycotans.[39]
Specialized organelle evolved from a mitochondrion, present in many anaerobicprotists. Like mitochondria, hydrogenosomes produce ATP and are involved in cellular respiration; in some cases, they still retain a genome.[10] They were discovered by Miklós Müller and Donald Lindmark in 1973 in the trichomonad genus Tritrichomonas.[52]
Plural incunabula. Elements added to the wall of a diatom auxospore before its expansion; they may include circular or elliptical scales, or narrow strips of silica.[26]
Also infusion animals. Organisms capable of producing dessication-resistant cysts (including some animals, like rotifers) which can be reactivated with an infusion of water. Coined by Martin Frobenius Ledermüller in the 18th century, it was later adopted as a zoological taxon by Jean Baptiste Lamarck. Since the acceptance of the cell theory until late in the 20th century, it was used exclusively as a synonym for ciliates.[22]
isogamous
The condition of performing the type of sexual reproduction known as anisogamy. Contrast: anisogamous.[5]
Also kathablepharid. A member of a group of phagotrophicflagellates closely related to cryptomonads. Their ejectisomes are composed of a single coiled ribbon, or "scroll", unlike the cryptomonads which have two scrolls per ejectisome.[17]
Refers to any fungus that lacks a flagellate stage in its life cycle. This includes aphelids, rozellids, microsporidians and chytrids. Although commonly regarded as fungi,[53] they were traditionally studied as protists to some degree.[22]
Disease caused by haemosporidian species (e.g., Plasmodium falciparum), spread by mosquitoes as vectors. Human malaria is one of the major serious diseases affecting humans, described more than 5,000 years ago and causing about half a million deaths annually.[23]
An organism with closed mitosis, absence of histones, and permanently condensed chromatin. The term was coined in 1965 by John David Dodge, who proposed that organisms with these characteristics represented an intermediate step in evolution between prokaryotes and "true" eukaryotes. Thus, according to this short-lived hypothesis, organisms like dinoflagellates had evolved earlier than the remaining eukaryotes.[55]
Also merogonial division. Phase in the life cycle of haemosporidians and coccidians where the meront goes through internal budding for asexual proliferation. The meronts bud into two or more daughter cells by cryptomitosis, and these cells develop into merozoites, which infect cells and differentiate again into meronts. Eventually, the last generation of meronts differentiate into macrogametocytes and microgametocytes instead, initiating the gamogony phase.[22][23]
merozoite
Stage in the life cycle of haemosporidians and coccidians produced by the meronts during merogony. The merozoites continue spreading the infection to other cells and becoming new meronts.[22][23]
Abbreviated MLO. Also mitochondrion-related organelle, abbreviated MRO. Term for organelles evolved from metabolically reduced mitochondria, namely hydrogenosomes and mitosomes.[10][58]
Organelle evolved from mitochondria that does not produce ATP or hydrogen, and lacks most metabolic pathways associated with mitochondria or hydrogenosomes.[10]
monad
Also -monas. Term used for nomenclatural purposes, mainly in reference to flagellates (see monadoid). Its initial use had a metaphysical meaning, to designate the indivisible and permanent smallest units of life, considered to be the elements and sources of all creatures. With the advent of optical microscopy and direct observation of unicellular life, the term was adapted by natural scientists during the 18-19th centuries to describe single-celled flagellates.[22]
Any member of Nucleariidae, a family of around 50 species of filosediscicristateamoebae that compose the closest relatives of fungi. Taxonomically, they belong to the monotypic order Rotosphaerida; consequently, they are sometimes called rotosphaerids. Some nucleariids are naked, while others are covered in scales. Among nucleariids is one species capable of aggregative multicellilarity, Fonticula alba.[39]
Stage in the life cycle of apicomplexans developed from the zygote and responsible for the production of sporocysts, which in turn produce sporozoites. This process known as sporogony. In some species, each oocyst only produces one sporocyst, making them equivalent.[23]
oogamous
The condition of performing the type of sexual reproduction known as oogamy.[49][5]
Any member of Piroplasmida, one of the two groups of hematozoans within the apicomplexans. Piroplasms include around 200 species of parasites of vertebrates that use ticks as vectors.[23]
In apusomonads, a highly mobile sleeve-like extension of folded membrane in the anterior region, which encloses the anterior flagellum; it is the primary characteristic of those organisms.[30]
Also protoctist. Any eukaryote that is not an animal (i.e., does not develop from a blastula), land plant (does not have embryonic stages)[61] or higher fungus (does not have a flagellate stage in their life cycles).[22]Lower fungi are also commonly excluded from protists.[6][53] The term is sometimes associated with any single-celled eukaryote instead, to the exclusion of macroalgae.[62] It was coined by Ernst Haeckel in 1866 as the taxon Protista, but under his definition it included prokaryotes and excluded ciliates. John Hogg introduced the taxon Protoctista in 1861 as a separate kingdom from those of animals, plants, and fungi. Under Robert H. Whittaker's 1959 concept of five kingdoms, the modern usage of protists as separate from prokaryotes and other eukaryotes was developed. [22]
Plural pseudopodia. Also pseudopod, with plural pseudopods. Extension of the cytoplasm produced in amoeboid organisms; may be extended or retracted for motility or to engulf food particles.[6]
Synonym of nucleariid. Due to the complicated taxonomic history of nucleariids, the first ones to be described were the naked, scale-less species under the family Nucleariidae; scale-bearing species (e.g., Pompholyxophrys) were described separately as Rotosphaerida by Heinrich Rainer in 1968. Later studies revealed that both belonged to the same group, and they are collectively known as nucleariids.[39]
Also slime mould. Informal (polyphyletic) category used to designate amoeboid organisms that, at some point in their life cycle, develop into a spore-bearing dispersal structure known as a fruting body.[6]
In the context of diatoms, specialized cell that undergoes a series of divisions to form sperm cells within itself;[26] equivalent to the antheridium in other protists.[21]
General term for specialized cells used for asexual dispersion or sexual reproduction; usually implies a degree of resistance to adverse conditions.[66]
Phase in the life cycle of apicomplexans where the zygote forms a protective wall around itself and differentiates into an oocyst, in which there are sporocysts that generate sporozoites, the infectious stage.[23]
Also tachyzoic merozoite. A merozoite characterized by rapid growth and replication, present in an early coccidial infection. Contrast: bradyzoite.[28]
Plural undulipodia. Motile appendage exclusive to eukaryotic cells. The term was popularized by Lynn Margulis to differentiate it from prokaryotic flagella; it is synonymous with flagella and cilia in the context of protists.[32][33][34]
unicell
Single-celled organism; the nominal counterpart of the adjective unicellular.[31]
Flagellate with a single flagellum associated to only one basal body.[71][6] The clade that groups together Amoebozoa and Opisthokonta was initially called Unikonta, due to a hypothesis where their common ancestor was a unikont; this was later refuted, and the clade name was changed to Amorphea.[71]
^ abSome single-celled protists, particularly flagellates such as choanoflagellates and many coccoid algae, are capable of forming colonies and may be labeled as multicellular in some instances.[39]
^Levandowsky, Michael (2012). "Physiological Adaptations of Protists". In Sperelakis, Nicholas (ed.). Cell Physiology Sourcebook: Essentials of Membrane Biophysics (Fourth ed.). Amsterdam; Boston: Elsevier/AP. pp. 874–890. ISBN978-0-12-387738-3.
^ abcSimpson, Alastair G. B.; Slamovits, Claudio H.; Archibald, John M. (2017). "Protist Diversity and Eukaryote Phylogeny". In Archibald, John M.; Simpson, Alastair G.B.; Slamovits, Claudio H. (eds.). Handbook of the Protists. Vol. 1 (2nd ed.). Springer. pp. 1–22. doi:10.1007/978-3-319-28149-0_45. ISBN978-3-319-28147-6.
^ abPánek, Tomáš; Simpson, Alastair G. B.; Brown, Matthew W.; Dyer, Betsey Dexter (2017). "Heterolobosea". In Archibald, John M.; Simpson, Alastair G.B.; Slamovits, Claudio H. (eds.). Handbook of the Protists. Vol. 2 (2nd ed.). Springer. pp. 1005–1046. doi:10.1007/978-3-319-28149-0_10. ISBN978-3-319-28147-6.
^ abHeiss AA, Lee WJ, Ishida KI, Simpson AG (2015). "Cultivation and Characterisation of New Species of Apusomonads (the Sister Group to Opisthokonts), Including Close Relatives of Thecamonas (Chelonemonas n. gen.)". Journal of Eukaryotic Microbiology. 62 (5): 637–649. doi:10.1111/jeu.12220. PMID25912654.
^ abcMcCourt, Richard M.; Karol, Kenneth G.; Hall, John D.; Casanova, Michelle T.; Grant, Michael C. (2017). "Charophyceae (Charales)". In Archibald, John M.; Simpson, Alastair G.B.; Slamovits, Claudio H. (eds.). Handbook of the Protists. Vol. 1 (2nd ed.). Springer. pp. 165–183. doi:10.1007/978-3-319-28149-0_40. ISBN978-3-319-28147-6.
^ abcdefMaistro, Silva; Broady, Paul; Andreoli, Carlo; Negrisolo, Enrico (2017). "Xanthophyceae". In Archibald, John M.; Simpson, Alastair G.B.; Slamovits, Claudio H. (eds.). Handbook of the Protists. Vol. 1 (2nd ed.). Springer. pp. 407–434. doi:10.1007/978-3-319-28149-0_30. ISBN978-3-319-28147-6.
^Torruella G, Galindo LJ, Moreira D, Ciobanu M, Heiss AA, Yubuki N, et al. (2022). "Expanding the molecular and morphological diversity of Apusomonadida, a deep-branching group of gliding bacterivorous protists". Journal of Eukaryotic Microbiology. 70 (2): e12956. doi:10.1111/jeu.12956. hdl:2117/404026. PMID36453005. S2CID253460648.
^ abcdefghMann, David G.; Crawford, Richard M.; Round, Frank E. (2017). "Bacillariophyta". In Archibald, John M.; Simpson, Alastair G.B.; Slamovits, Claudio H. (eds.). Handbook of the Protists. Vol. 1 (2nd ed.). Springer. pp. 205–266. doi:10.1007/978-3-319-28149-0_29. ISBN978-3-319-28147-6.
^ abcBeakes, Gordon W.; Thines, Marco (2017). "Hyphochytriomycota and Oomycot". In Archibald, John M.; Simpson, Alastair G.B.; Slamovits, Claudio H. (eds.). Handbook of the Protists. Vol. 1 (2nd ed.). Springer. pp. 435–505. doi:10.1007/978-3-319-28149-0_26. ISBN978-3-319-28147-6.
^ abcBerney, Cédric; Geisen, Stefan; Van Wichelen, Jeroen; Nitsche, Frank; Vanormelingen, Pieter; Bonkowski, Michael; Bass, David (20 April 2015). "Expansion of the 'Reticulosphere': Diversity of Novel Branching and Network-forming Amoebae Helps to Define Variosea (Amoebozoa)". Protist. 166 (2): 271–295. doi:10.1016/j.protis.2015.04.001. PMID25965302.
^ abcdHeiss, Aaron A.; Brown, Matthew W.; Simpson, Alastair G. B. (2017). "Apusomonadida". In Archibald, John M.; Simpson, Alastair G.B.; Slamovits, Claudio H. (eds.). Handbook of the Protists. Vol. 1 (2nd ed.). Springer. p. 1619–1645. doi:10.1007/978-3-319-28149-0_15. ISBN978-3-319-28147-6.
^ abcdeLeander, Brian S.; Lax, Gordon; Karnkowska, Anna; Simpson, Alastair G. B. (2017). "Euglenida". In Archibald, John M.; Simpson, Alastair G.B.; Slamovits, Claudio H. (eds.). Handbook of the Protists. Vol. 2 (2nd ed.). Springer. pp. 1047–1088. doi:10.1007/978-3-319-28149-0_13. ISBN978-3-319-28147-6.
^ abcAndersen, R. A.; Barr, D. J. S.; Lynn, D. H.; Melkonian, M.; Moestrup, Ø.; Sleigh, M. A. (1991). "Terminology and nomenclature of the cytoskeletal elements associated with the flagellar/ciliary apparatus in protists". Protoplasma. 164 (1–3): 1–8. Bibcode:1991Prpls.164....1A. doi:10.1007/BF01320809.
^ abcdePrice, Dana C.; Steiner, Jürgen M.; Yoon, Hwan Su; Bhattacharya, Debashish; Löffelhardt, Wolfgang (2017). "Glaucophyta". In Archibald, John M.; Simpson, Alastair G.B.; Slamovits, Claudio H. (eds.). Handbook of the Protists. Vol. 1 (2nd ed.). Springer. pp. 23–87. doi:10.1007/978-3-319-28149-0_42. ISBN978-3-319-28147-6.
^ abBrown MW, et al. (2018), "Phylogenomics Places Orphan Protistan Lineages in a Novel Eukaryotic Super-Group", Genome Biology and Evolution, 10 (2): 427–433, doi:10.1093/gbe/evy014, PMC5793813, PMID29360967
^ abSimpson, Alastair G. B. (2017). "Jakobida". In Archibald, John M.; Simpson, Alastair G.B.; Slamovits, Claudio H. (eds.). Handbook of the Protists. Vol. 2 (2nd ed.). Springer. pp. 973–1004. doi:10.1007/978-3-319-28149-0_6. ISBN978-3-319-28147-6.
^ abcBass, David; Chao, Ema E.-Y.; Nikolaev, Sergey; Yabuki, Akinori; Ishida, Ken-Ichiro; Berney, Cédric; Pakzad, Ursula; Wylezich, Claudia; Cavalier-Smith, Thomas (7 October 2008). "Phylogeny of novel naked filose and reticulose Cercozoa: Granofilosea cl. n. and Proteomyxidea revised". Protist. 160: 75–109. doi:10.1016/j.protis.2008.07.002.
^ abBrown, Matthew W.; Silberman, Jeffrey D.; Spiegel, Frederick W. (26 November 2010). ""Slime Molds" among the Tubulinea (Amoebozoa): Molecular Systematics and Taxonomy of Copromyxa". Protist. 162 (2): 277–287. doi:10.1016/j.protis.2010.09.003. PMID21112814.
^ abcKawai, Hiroshi; Henry, Eric C. (2017). "Phaeophyta". In Archibald, John M.; Simpson, Alastair G.B.; Slamovits, Claudio H. (eds.). Handbook of the Protists. Vol. 1 (2nd ed.). Springer. pp. 267–304. doi:10.1007/978-3-319-28149-0_31. ISBN978-3-319-28147-6.
^ abcdPage, Frederick C. (1987). "The classification of 'naked' amoebae (Phylum Rhizopoda)". Archiv für Protistenkunde. 133 (3–4): 199–217. doi:10.1016/S0003-9365(87)80053-2.
^ abTedersoo, Leho; Sánchez-Ramírez, Santiago; Kõljalg, Urmas; Bahram, Mohammad; Döring, Markus; Schigel, Dmitry; May, Tom; Ryberg, Martin; Abarenkov, Kessy (2018), "High-level classification of the Fungi and a tool for evolutionary ecological analyses", Fungal Diversity, 90: 135–159, doi:10.1007/s13225-018-0401-0, hdl:10138/238983, S2CID21714270
^ abKostka, Martin (2017). "Opalinata". In Archibald, John M.; Simpson, Alastair G.B.; Slamovits, Claudio H. (eds.). Handbook of the Protists. Vol. 1 (2nd ed.). Springer. pp. 543–565. doi:10.1007/978-3-319-28149-0_4. ISBN978-3-319-28147-6.
^ abTaylor, F. J. R. (February 1976). "Flagellate Phylogeny: A Study in Conflicts". The Journal of Protozoology. 23 (1): 28–40. doi:10.1111/j.1550-7408.1976.tb05242.x.
^O'Malley MA, Simpson AG, Roger AJ (2012). "The other eukaryotes in light of evolutionary protistology". Biology & Philosophy. 28 (2): 299–330. doi:10.1007/s10539-012-9354-y. S2CID85406712.
^ abcSpiegel, Frederick W.; Shadwick, Lora L.; Ndiritu, George G.; Brown, Matthew W.; Aguilar, Maria; Shadwick, John D. (2017). "Protosteloid amoebae (Protosteliida, Protosporangiida, Cavosteliida, Schizoplasmodiida, Fractoviteliida, and sporocarpic members of Vannellida, Centramoebida, and Pellitida)". In Archibald, John M.; Simpson, Alastair G.B.; Slamovits, Claudio H. (eds.). Handbook of the Protists. Vol. 2 (2nd ed.). Springer. doi:10.1007/978-3-319-28149-0_12. ISBN978-3-319-28147-6.
^Jeon, Boo Seong; Park, Myung Gil (27 June 2020). "Parvilucifera multicavata sp. nov. (Alveolata, Perkinsozoa), a new parasitoid infecting marine dinoflagellates having abundant apertures on the sporangium". Protist. 171: 125743. doi:10.1016/j.protis.2020.125743.
^Saldarriaga, Juan F.; Taylor, F. J. R. 'Max' (2017). "Dinoflagellata". In Archibald, John M.; Simpson, Alastair G.B.; Slamovits, Claudio H. (eds.). Handbook of the Protists. Vol. 1 (2nd ed.). Springer. pp. 625–678. doi:10.1007/978-3-319-28149-0_22. ISBN978-3-319-28147-6.
^Tice, Alexander K.; Spiegel, Frederick W.; Brown, Matthew W. (24 February 2023). "Phylogenetic placement of the protosteloid amoeba Microglomus paxillus identifies another case of sporocarpic fruiting in Discosea (Amoebozoa)". Journal of Eukaryotic Microbiology. 70 (4): e12971. doi:10.1111/jeu.12971.
^Leander, Brian S.; Lax, Gordon; Karnkowska, Anna; Simpson, Alastair G. B. (2017). "Euglenida". In Archibald, John M.; Simpson, Alastair G.B.; Slamovits, Claudio H. (eds.). Handbook of the Protists. Vol. 2 (2nd ed.). Springer. pp. 1047–1088. doi:10.1007/978-3-319-28149-0_13. ISBN978-3-319-28147-6.
^ abAdl, Sina M.; Simpson, Alastair G. B.; Lane, Christopher E.; Lukeš, Julius; Bass, David; Bowser, Samuel S.; Brown, Matthew W.; Burki, Fabien; Dunthorn, Micah; Hampl, Vladimir; Heiss, Aaron; Hoppenrath, Mona; Lara, Enrique; le Gall, Line; Lynn, Denis H.; McManus, Hilary; Mitchell, Edward A. D.; Mozley-Stanridge, Sharon E.; Parfrey, Laura W.; Pawlowski, Jan; Rueckert, Sonja; Shadwick, Laura; Schoch, Conrad L.; Smirnov, Alexey; Spiegel, Frederick W. (28 September 2012). "The Revised Classification of Eukaryotes". The Journal of Eukaryotic Microbiology. 59 (2): 429–514. doi:10.1111/j.1550-7408.2012.00644.x. PMC3483872. PMID23020233.
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