Superregnum: Eukaryota
Supergroup: Unikonta
Cladus: Opisthokonta
Cladi: Animalia (regnum) - Choanomonada - Fungi (regnum) - Mesomycetozoa
Name
Opisthokonta Cavalier-Smith 1987, emend. Adl et al. 2005
Alternative classifications
Vischer (1945)
Vischer, W. (1945). Über einen pilzähnlichen, autotrophen Mikroorganismus, Chlorochytridion, einige neue Protococcales und die systematische Bedeutung der Chloroplasten. Verhandlungen der Naturforschenden Gesellschaft in Basel 6: 41–49. From Leadbeater (2015), [1].
Opistokontae (Pulselloflagellatae)
- choanoflagellates
- chytrids
- ”some uniflagellate algal monads”
Gams (1947)
Gams, H. (1947). Die Protochlorinae als autotrophe Vorfahren von Pilzen und Tieren? Mikroskopie, 2, 383–7, [2]. From Leadbeater (2015) and Copeland (1956).
Opistokonten
- choanoflagellates
- chytrids
- ”some uniflagellate algal monads” [Protochlorinae? Pedilomonas, Chlorochytridion?]
- metazoans
- sporozoans
Rothmaler (1951)
Rothmaler, Werner. (1951). Die Abteilungen und Klassen der Pflanzen. Feddes Rep Spec Nov Reg Veg 3: 256–266, [3], [4].
- Protobionta Rothm. (1948) – Protozoa Owen (1860) – Protista Haeckel (1866) emend. Barkley (1939) – Protoctista Hogg (1861)
- Mycophyta Pascher (1931) – Inophyta Haeckel (1866) – Phaeophyta Mycophytina Rothm. (1949) [?]
- Archimycota Rothm., nom. nov. – Archimycetes A. Fischer (1894) – Opisthocontae Gams (1947) – Protochlorinae Vischer (1945) – Opisthophyceae Rothm., nom. event. – Archemycetae Barkley (1939)
- Mycophyta Pascher (1931) – Inophyta Haeckel (1866) – Phaeophyta Mycophytina Rothm. (1949) [?]
Copeland (1956)
Copeland, H. F. (1956). The Classification of Lower Organisms. Palo Alto: Pacific Books, [5].
- Kingdom Protoctista
- Phylum Opisthokonta
- Class Archimycetes
- Order 1. Monoblepharidalea
- Order 2. Chytridinea
- Class Archimycetes
- Phylum Opisthokonta
Cavalier-Smith (1987)
Cavalier-Smith, T. (1987). The origin of fungi and pseudofungi. In: Rayner, Alan D. M. (ed.). Evolutionary biology of Fungi. Cambridge: Cambridge Univ. Press. pp. 339–353. From Leadbeater (2015).
Opisthokonta
- choanoflagellates
- animals
- fungi
Hausmann et al. (2003)
From Hausmann, K., N. Hulsmann, R. Radek. Protistology. Schweizerbart'sche Verlagsbuchshandlung, Stuttgart, 2003.
Empire Eukaryota Chatton, 1925 (= Eukarya)
- Phylum Opisthokonta Cavalier-Smith & Chao, 1995
- Subphylum Fungi Nees,1817
- Infraphylum Chytridiomycota de Вагу, 1863
- Infraphylum Zygomycota Barr, 1982
- Infraphylum Eumycota Cavalier-Smith, 1998 (Dikaryomycota)
- Superclass Microspora Sprague, 1982
- Class Microsporea Sprague, 1982
- Subclass Rudimicrosporia Sprague, 1977
- Subclass Microsporia Delphy, 1963
- Class Microsporea Sprague, 1982
- Superclass Ascomycota Berkeley, 1857
- Class Archaeascomycota
- Class Hemiascomycota
- Class Euascomycota
- Family Nephridiophagidae Sprague, 1970
- Superclass Basidiomycota de Вагу, 1866
- Superclass Microspora Sprague, 1982
- Subphylum Choanozoa Cavalier-Smith, 1981
- Superclass Mesomycetozoa Herr et al., 1999
- Superclass Choanoflagellata Kent, 1880
- Infraphylum Metazoa Haeckel, 1874
- Superclass Myxozoa Grassé, 1970
- Subphylum Fungi Nees,1817
Adl et al. (2012)
Adl, S.M. et al. (2012). The revised classification of eukaryotes. Journal of Eukaryotic Microbiology 59 (5): 429–514, [6].
Eukaryota
- Opisthokonta Cavalier-Smith 1987, emend. Adl et al. 2005
- Holozoa Lang et al. 2002 (R)
- Filasterea Shalchian-Tabrizi et al. 2008
- Ministeria Patterson et al. 1993, emend. Tong 1997 (Ministeriida Cavalier-Smith, 1997)
- Capsaspora Hertel et al. 2002 (Capsasporidae Cavalier-Smith 2008) (M)
- Ichthyosporea Cavalier-Smith 1998 (Mesomycetozoea Mendoza et al. 2002)
- Rhinosporidaceae Mendoza et al. 2001 (Dermocystida Cavalier-Smith 1998) (R)
- Amphibiocystidium ranae, Amphibiothecum penneri, Dermocystidium, Rhinosporidium seeberi, Sphaerothecum destruens
- Ichthyophonae Mendoza et al. 2001 (Ichthyophonida Cavalier-Smith 1998; Amoebidiidae Reeves 2003) (R)
- Abeoforma whisleri, Amoebidium parasiticum, Anurofeca richardsi, Astreptonema, Caullerya mesnili, Creolimax fragrantissima, Eccrinidus flexilis, Enterobryus oxidi, Enteropogon sexuale, Ichthyophonus, Palavascia patagonica, Pseudoperkinsus tapetis, Psorospermium haeckeli, Sphaeroforma arctica
- Rhinosporidaceae Mendoza et al. 2001 (Dermocystida Cavalier-Smith 1998) (R)
- Aphelidea Gromov 2000
- Amoeboaphelidium, Aphelidium, Pseudoaphelidium [Pseudaphelidium]
- Corallochytrium Raghu-Kumar 1987 (M)
- Choanomonada Kent 1880
- Craspedida Cavalier-Smith 1997, emend. Nitsche et al. 2011
- Astrosiga, Aulomonas, Choanoeca, Cladospongia, Codonocladium, Codonosigopsis, Codosiga (junior synonym Codonosiga), Desmarella (junior synonyms Codonodesmus and Kentrosiga), Dicraspedella, Diploeca, Diplosiga, Diplosigopsis, Kentia, Lagenoeca, Monosiga, Pachysoeca, Proterospongia, Salpingoeca, Salpingorhiza, Sphaeroeca, Stelexomonas, Stylochromonas
- Acanthoecida Norris 1965, emend. Cavalier-Smith 1997, emend. Nitsche et al. 2011
- Acanthoeca, Acanthocorbis, Amoenoscopa, Apheloecion, Bicosta, Calliacantha, Calotheca, Campanoeca, Campyloacantha, Conion, Cosmoeca, Crinolina, Crucispina, Diaphanoeca, Didymoeca, Helgoeca, Kakoeca, Monocosta, Nannoeca, Parvicorbicula, Platypleura, Pleurasiga, Polyfibula, Polyoeca, Saepicula, Saroeca, Savillea, Spinoeca, Spiraloecion, Stephanacantha, Stephanoeca, Syndetophyllum
- Craspedida Cavalier-Smith 1997, emend. Nitsche et al. 2011
- Metazoa Haeckel 1874
- Porifera Grant 1836 (Parazoa Sollas 1884)
- Silicispongia Schmidt 1862 (Silicea Bowerbank 1864, emend. Gray 1867)
- Hexactinellida Schmidt 1870
- Demospongiae Sollas 1885, emend. Borchiellini et al. 2004
- Democlavia Sperling et al. 2009, emend. Morrow et al. 2012 (R)
- Homoscleromorpha Lévi 1973, emend. Borchiellini et al. 2004 (R)
- Calcispongia Johnston 1842 (Calcarea Bowerbank 1864)
- Calcinea Hartman 1958, emend. Borchiellini et al. 2004 (R)
- Calcaronea Hartman 1958, emend. Borchiellini et al. 2004 (R)
- Silicispongia Schmidt 1862 (Silicea Bowerbank 1864, emend. Gray 1867)
- Trichoplax von Schulze 1883 (Placozoa Grell 1971) (M)
- Animalia Linnaeus 1758, emend. Adl et al. 2005 (Eumetazoa Bütschli 1910) [including Myxozoa Grassé 1970]
- Porifera Grant 1836 (Parazoa Sollas 1884)
- Filasterea Shalchian-Tabrizi et al. 2008
- Nucletmycea Brown et al. 2009 (Holomycota Liu et al. 2009) (R)
- Nuclearia Cienkowski 1865
- Fonticula Worley et al. 1979 (M)
- Rozella Cornu 1872 (= Rozellida Lara et al. 2010; Cryptomycota M. D. M. Jones & T. A. Richards 2011)
- Fungi R. T. Moore 1980
- Holozoa Lang et al. 2002 (R)
[Notes: M, monotypic group with only one described species; P, paraphyletic group; R, ribogroup assembled from phylogenetic studies.]
Karpov et al. (2014)
Karpov, S. A., Mamkaeva, M. A., Aleoshin, V. V., Nassonova, E., Lilje, O., & Gleason, F. H. (2014). Morphology, phylogeny, and ecology of the aphelids (Aphelidea, Opisthokonta) and proposal for the new superphylum Opisthosporidia. Frontiers in Microbiology, 5: 112. doi: 10.3389/fmicb.2014.00112.
Opisthokonta Cavalier-Smith, 1987
- Holozoa
- Metazoa
- Choanoflagellata
- Mesomycetozoea
- Holomycota
- Nucleariida
- Fungi
- Opisthosporidia Karpov, Aleoshin et Mikhailov superphylum nov.
- Cryptomycota (Jones et Richards, 2011), emend. Karpov et Aleoshin
- Aphelida Karpov, Aleoshin et Mikhailov phylum nov.
- Aphelidea Gromov, 2000
- Microsporidia
Ruggiero et al. (2015)
Ruggiero, M.A., Gordon, D.P., Orrell, T.M., Bailly, N., Bourgoin, T., Brusca, R.C., Cavalier-Smith, T., Guiry, M. D. & Kirk, P. M. (2015). A Higher Level Classification of All Living Organisms. PLoS ONE 10(4): e0119248, [7].
- Superkingdom Prokaryota
- Kingdom Archaea [= Archaebacteria]
- Kingdom Bacteria [= Eubacteria]
- Superkingdom Eukaryota
- Kingdom Protozoa
- Subkingdom Eozoa
- Infrakingdom Euglenozoa
- Infrakingdom Excavata
- Subkingdom Sarcomastigota
- Phylum Amoebozoa
- Phylum Choanozoa [with Microsporidia, Animalia, and Fungi constitutes "Supergroup Opisthokonta"]
- Phylum Microsporidia [with Choanozoa, Animalia, and Fungi constitutes "Supergroup Opisthokonta"]
- Phylum Sulcozoa
- Subkingdom Eozoa
- Kingdom Chromista
- Subkingdom Hacrobia
- Subkingdom Harosa [= "Supergroup SAR"]
- Infrakingdom Halvaria
- Superphylum Alveolata
- Superphylum Heterokonta [= "Supergroup Stramenopiles"]
- Infrakingdom Rhizaria
- Infrakingdom Halvaria
- Kingdom Fungi
- Subkingdom Dikarya [= Neomycota]
- Subkingdom Eomycota
- Kingdom Plantae [= Archaeplastida]
- Subkingdom Biliphyta
- Subkingdom Viridiplantae
- Kingdom Animalia
- Kingdom Protozoa
Toruella et al. (2015)
Torruella, Guifré, et al. Phylogenomics reveals convergent evolution of lifestyles in close relatives of animals and fungi. Current Biology 25: 1–7, [8].
- Holozoa
- Metazoa
- Choanoflagellatea
- Filasterea
- "Teretosporea"
- Ichthyosporea
- Ichthyophonida
- Sphaerothecum
- Corallochytrium
- Ichthyosporea
- Holomycota
- Discicristoidea
- Fonticula
- Nuclearia
- Microsporidia
- Cryptomycota
- Rozella
- Fungi
- Discicristoidea
References
- Cavalier-Smith, T. 2009: Megaphylogeny, cell body plans, adaptive zones: causes and timing of eukaryote basal radiations. Journal of eukaryotic microbiology, 56: 26–33. doi: 10.1111/j.1550-7408.2008.00373.x
- Philip, G.K.; Creevey, C.J.; McInerney, J.O. 2005: The Opisthokonta and the Ecdysozoa May Not Be Clades: Stronger Support for the Grouping of Plant and Animal than for Animal and Fungi and Stronger Support for the Coelomata than Ecdysozoa. Molecular biology and evolution, 22: 1175–1184. doi: 10.1093/molbev/msi102
- Steenkamp, E.T.; Wright, J.; Baldauf, S.L. 2006: The protistan origins of animals and fungi. Molecular biology and evolution, 23: 93–106. doi: 10.1093/molbev/msj011
links
- Prous, Marko. The Opisthokonta and the Ecdysozoa may be clades: An update on Philip et al. (2005). Available from Nature Precedings doi: 10.1038/npre.2009.3306.1 (2009)
- Wegener Parfrey, Laura 2009. Taxon page for Eukaryota. In: The ATOL eutree hypothesis First Created: 2009–07-04T15:11:59Z. Last Updated: 2009–07-04T15:14:20Z.
Vernacular names
Ελληνικά: ΟπισθόκονταEnglish: Opisthokont
français: Opisthocontes, opisthocontes
македонски: Заднокамшични
日本語: オピストコンタ
Tiếng Việt: Sinh vật lông roi sau
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The opisthokonts (Greek: ὀπίσθιος (opísthios) = "rear, posterior" + κοντός (kontós) = "pole" i.e. "flagellum") or Choanozoa are a broad group of eukaryotes, including both the animal and fungus kingdoms,[5] together with the eukaryotic microorganisms that are sometimes grouped in the paraphyletic phylum Choanozoa (conventionally assigned to the protist "kingdom").[6] The opisthokonts, sometimes referred to as the "Fungi/Metazoa group",[7] are generally recognized as a monophyletic clade, and it is thought to be a sister to the Apusomonadida clade.[6][8][9][10]
Flagella
A common characteristic of opisthokonts is that flagellate cells, such as the sperm of most animals and the spores of the chytrid fungi, propel themselves with a single posterior flagellum. It is this feature that gives the group its name. In contrast, flagellate cells in other eukaryote groups propel themselves with one or more anterior flagella. However, in some opisthokont groups, including most of the fungi, flagellate cells have been lost.[6]
History
The close relationship between animals and fungi was suggested by Thomas Cavalier-Smith in 1987,[3] who used the informal name opisthokonta (the formal name has been used for the chytrids by Copeland in 1956), and was supported by later genetic studies.[11]
Early phylogenies placed fungi near the plants and other groups that have mitochondria with flat cristae, but this character varies. More recently, it has been said that holozoa (animals) and holomycota (fungi) are much more closely related to each other than either is to plants, because opisthokonts have a triple fusion of carbamoyl phosphate synthetase, dihydroorotase, and aspartate carbamoyltransferase that is not present in plants, and plants have a fusion of thymidylate synthase and dihydrofolate reductase not present in the opisthokonts. Animals and fungi are also more closely related to amoebas than they are to plants, and plants are more closely related to the SAR supergroup of protists than they are to animals or fungi.[citation needed] Animals and fungi are both heterotrophs, unlike plants, and while fungi are sessile like plants, there are also sessile animals.
Cavalier-Smith and Stechmann[12] argue that the uniciliate eukaryotes such as opisthokonts and Amoebozoa, collectively called unikonts, split off from the other biciliate eukaryotes, called bikonts, shortly after they evolved.
Taxonomy
Opisthokonts are divided into Holomycota or Nucletmycea (fungi and all organisms more closely related to fungi than to animals) and Holozoa (animals and all organisms more closely related to animals than to fungi); no opisthokonts basal to the Holomycota/Holozoa split have yet been identified.[citation needed] The Opisthokonts was largely resolved by Torriella et al.[13]. Holomycota and Holozoa are composed of the following groups.
- Holomycota
- Fungi
- including:
- chytrids (flagellated, zoosporic fungi)
- microsporidia (previously thought to be sporozoans)
- Hyaloraphidium (previously thought to be a green alga, now considered a chytrid)
- excluding:
- oomycetes (water molds) (now included in stramenopiles)
- labyrinthulomycetes (slime nets) (now included in stramenopiles)
- myxomycetes (now included in amoebozoans)
- including:
- Rozellida (previously considered chytrids)
- Fonticula (a slime mold, formerly considered an acrasiomycete)
- Nucleariida or Cristidiscoidea (nucleariids, filose amoebae)
- Fungi
- Holozoa
- Mesomycetozoea
- Dermocystida (formerly considered parasitic fungi or sporozoans)
- Ichthyophonida (formerly considered parasitic fungi incertae sedis)
- Eccrinales (formerly considered trichomycetes)
- Amoebidiales (formerly considered trichomycetes)
- Corallochytrium (formerly considered a thraustochytrid)
- Filozoa
- Filasterea
- Capsasporida (capsasporids, newly discovered amoebae)
- Ministeriida (ministeriids, newly discovered amoebae)
- Choanoflagellata (flagellates formerly included in protozoa)
- Animalia
- including Myxozoa (previously considered sporozoans, now considered derived from cnidarians)
- Filasterea
- Mesomycetozoea
The paraphyletic taxon Choanozoa includes either non-animal holozoans, or non-animal, non-fungi opisthokonts.
The choanoflagellates have a circular mitochondrial DNA genome with long intergenic regions. This is four times as large as animal mitochondrial genomes and contains twice as many protein genes.
Corallochytrium seem likely to be more closely related to the fungi than to the animals on the basis of the presence of ergosterol in their membranes and being capable of synthesis of lysine via the α-aminoadipate (AAA) pathway.
The ichthyosporeans have a two amino acid deletion in their elongation factor 1 α gene that is considered characteristic of fungi.
The ichthyosl genome is >200 kilobase pairs in length and consists of several hundred linear chromosomes that share elaborate terminal-specific sequence patterns.
Eukaryotes |
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One view of the great kingdoms and their stem groups.[14]
References
- Eichinger, L.; Pachebat, J. A.; Glöckner, G.; Rajandream, M. A.; Sucgang, R.; Berriman, M.; Song, J.; Olsen, R.; et al. (2005). "The genome of the social amoeba Dictyostelium discoideum". Nature. 435 (7038): 43–57. PMC 1352341 . PMID 15875012. doi:10.1038/nature03481.
- Steenkamp, E. T.; Wright, J.; Baldauf, S. L. (2005). "The Protistan Origins of Animals and Fungi". Molecular Biology and Evolution. 23 (1): 93–106. PMID 16151185. doi:10.1093/molbev/msj011.
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