Euglenozoa Explained
Euglenozoa are a large group of flagellate Discoba. They include a variety of common free-living species, as well as a few important parasites, some of which infect humans. Euglenozoa are represented by four major groups, i.e., Kinetoplastea, Diplonemea, Euglenida, and Symbiontida. Euglenozoa are unicellular, mostly around NaNμm in size, although some euglenids get up to 500μm long.[1]
Structure
Most euglenozoa have two flagella, which are inserted parallel to one another in an apical or subapical pocket. In some these are associated with a cytostome or mouth, used to ingest bacteria or other small organisms. This is supported by one of three sets of microtubules that arise from the flagellar bases; the other two support the dorsal and ventral surfaces of the cell.[2]
Some other euglenozoa feed through absorption, and many euglenids possess chloroplasts, the only eukaryotes outside Diaphoretickes to do so without performing kleptoplasty,[3] and so obtain energy through photosynthesis. These chloroplasts are surrounded by three membranes and contain chlorophylls A and B, along with other pigments, so are probably derived from a green alga, captured long ago in an endosymbiosis by a basal euglenozoan. Reproduction occurs exclusively through cell division. During mitosis, the nuclear membrane remains intact, and the spindle microtubules form inside of it.[2]
The group is characterized by the ultrastructure of the flagella. In addition to the normal supporting microtubules or axoneme, each contains a rod (called paraxonemal), which has a tubular structure in one flagellum and a latticed structure in the other. Based on this, two smaller groups have been included here: the diplonemids and Postgaardi.[4]
Classification
Historically, euglenozoans have been treated as either plants or animals, depending on whether they belong to largely photosynthetic groups or not. Hence they have names based on either the International Code of Nomenclature for algae, fungi, and plants (ICNafp) or the International Code of Zoological Nomenclature (ICZN). For example, one family has the name Euglenaceae under the ICNafp and the name Euglenidae under the ICZN. As another example, the genus name Dinema is acceptable under the ICZN, but illegitimate under the ICNafp, as it is a later homonym of an orchid genus, so that the synonym Dinematomonas must be used instead.
The Euglenozoa are generally accepted as monophyletic. They are related to Percolozoa; the two share mitochondria with disk-shaped cristae, which only occurs in a few other groups.[5] Both probably belong to a larger group of eukaryotes called the Excavata.[6] This grouping, though, has been challenged.[7]
Phylogeny
The phylogeny based on the work of Cavalier-Smith (2016):[8]
A consensus phylogeny following the review by Kostygov et al. (2021):[9]
Taxonomy
Cavalier-Smith (2016/2017)
The following classification of Euglenozoa is as described by Cavalier-Smith in 2016, modified to include the new subphylum Plicomonada according to Cavalier-Smith et al (2017).[10]
Phylum Euglenozoa Cavalier-Smith 1981 emend. Simpson 1997 [Euglenobionta]
- Subphylum Glycomonada Cavalier-Smith 2016
- Class Diplonemea Cavalier-Smith 1993 emend. Simpson 1997 [Diplosonematea; Diplonemia <small>Cavalier-Smith 1993</small>]
- Order Diplonemida Cavalier-Smith 1993 [Hemistasiida]
- Family Hemistasiidae Cavalier-Smith 2016 [Entomosigmaceae]
- Family Diplonemidae Cavalier-Smith 1993 [Rhynchopodaceae <small>Skuja 1948 ex Cavalier-Smith 1993</small>]
- Class Kinetoplastea Honigberg 1963 emend. Margulis 1974 [Kinetoplastida <small>Honigberg 1963</small>; Kinetoplasta <small>Honigberg 1963 stat. nov.</small>]
- Subclass Prokinetoplastina Vickerman 2004
- Order Prokinetoplastida Vickerman 2004
- Family Ichthyobodonidae Isaksen et al., 2007
- Subclass Metakinetoplastina Vickerman 2004
- Order Bodonida* Hollande 1952 em. Vickerman 1976, Kryov et al 1980
- Suborder Neobodonida Vickerman 2004
- Family Rhynchomonadidae Cavalier-Smith 2016
- Family Neobodonidae Cavalier-Smith 2016
- Suborder Parabodonida Vickerman 2004
- Family Parabodonidae Cavalier-Smith 2016
- Family Cryptobiidae* Vickerman 2004
- Suborder Eubodonida Vickerman 2004
- Family Bodonidae Bütschli 1883 [Bodonaceae <small>Lemmermann 1914</small>; Bodoninae <small>Bütschli 1883</small>; Pleuromonadidae <small>Kent 1880</small>]
- Order Trypanosomatida Kent 1880 stat. n. Hollande, 1952 emend. Vickerman 2004
- Subphylum Plicomonada Cavalier-Smith 2017
- Infraphylum Postgaardia Cavalier-Smith 2016 stat. nov. Cavalier-Smith 2017
- Class Postgaardea Cavalier-Smith 1998 s.s. [Symbiontida <small>Yubuki et al., 2009</small>]
- Order Bihospitida Cavalier-Smith 2016
- Family Bihospitidae Cavalier-Smith 2016
- Order Postgaardida Cavalier-Smith 2003
- Infraphylum Euglenoida Bütschli 1884 emend. Senn 1900 stat. nov. Cavalier-Smith, 2017 [Euglenophyta; Euglenida <small>Buetschli 1884</small>; Euglenoidina <small>Buetschli 1884</small>]
- Parvphylum Entosiphona Cavalier-Smith 2016 stat. nov. Cavalier-Smith 2017
- Class Entosiphonea Cavalier-Smith 2016
- Order Entosiphonida Cavalier-Smith 2016
- Family Entosiphonidae Cavalier-Smith 2016
- Parvphylum Dipilida Cavalier-Smith 2016 stat. nov. Cavalier-Smith 2017
- Superclass Rigimonada* Cavalier-Smith 2016
- Class Stavomonadea Cavalier-Smith 2016 [Petalomonadea <small>Cavalier-Smith 1993</small>; Petalomonadophyceae]
- Subclass Heterostavia Cavalier-Smith 2016
- Order Heterostavida Cavalier-Smith 2016
- Family Serpenomonadidae Cavalier-Smith 2016
- Subclass Homostavia Cavalier-Smith 2016
- Order Decastavida Cavalier-Smith 2016a
- Family Decastavidae Cavalier-Smith 2016a
- Family Keelungiidae Cavalier-Smith 2016a
- Order Petalomonadida Cavalier-Smith 1993 [Sphenomonadales <small>Leedale 1967</small>; Sphenomonadina <small>Leedale 1967</small>]
- Family Sphenomonadidae Kent 1880
- Family Petalomonadidae [Petalomonadaceae <small>Buetschli 1884</small>; Notosolenaceae <small>Stokes 1888</small>; Scytomonadaceae <small>Ritter von Stein 1878</small>]
- Class Ploeotarea Cavalier-Smith 2016
- Order Ploeotiida Cavalier-Smith 1993
- Family Lentomonadidae Cavalier-Smith 2016
- Family Ploeotiidae Cavalier-Smith 2016
- Superclass Spirocuta Cavalier-Smith 2016
- Class Peranemea Cavalier-Smith 1993 emend. Cavalier-Smith 2016
- Subclass Acroglissia Cavalier-Smith 2016
- Order Acroglissida Cavalier-Smith 2016
- Family Teloproctidae Cavalier-Smith 2016a
- Subclass Peranemia Cavalier-Smith 2016
- Order Peranemida Bütschli 1884 stat. nov. Cavalier-Smith 1993
- Family Peranematidae [Peranemataceae <small>Dujardin 1841</small>; Pseudoperanemataceae <small>Christen 1962</small>]
- Subclass Anisonemia Cavalier-Smith 2016
- Order Anisonemida Cavalier-Smith 2016 [Heteronematales <small>Leedale 1967</small>]
- Family Anisonemidae Saville Kent, 1880 em. Cavalier-Smith 2016 [Heteronemidae <small>Calkins 1926</small>; Zygoselmidaceae <small>Kent 188</small>]
- Order Natomonadida Cavalier-Smith 2016
- Suborder Metanemina Cavalier-Smith 2016
- Suborder Rhabdomonadina Leedale 1967 emend. Cavalier-Smith 1993 [Astasida <small>Ehrenberg 1831</small>; Rhabdomonadia <small>Cavalier-Smith 1993</small>; Rhabdomonadophyceae; Rhabdomonadales]
- Family Distigmidae Hollande, 1942
- Family Astasiidae Saville Kent, 1884 [Astasiaceae <small>Ehrenberg orth. mut. Senn 1900</small>; Rhabdomonadaceae <small>Fott 1971</small>; Menoidiaceae <small>Buetschli 188</small>; Menoidiidae <small>Hollande, 1942</small>]
- Class Euglenophyceae Schoenichen 1925 emend. Marin & Melkonian 2003 [Euglenea <small>Bütschli 1884 emend. Busse & Preisfeld 2002</small>; Euglenoidea <small>Bütschli 1884</small>; Euglenida <small>Bütschli 1884</small>] (Photosynthetic clade)
- Subclass Rapazia Cavalier-Smith 2016
- Subclass Euglenophycidae Busse and Preisfeld, 2003
- Order Eutreptiida [Eutreptiales <small>Leedale 1967 emend. Marin & Melkonian 2003</small>; Eutreptiina <small>Leedale 1967</small>]
- Family Eutreptiaceae [Eutreptiaceae <small>Hollande 1942</small>]
- Order Euglenida Ritter von Stein, 1878 stat. n. Calkins, 1926 [Euglenales <small>Engler 1898 emend. Marin & Melkonian 2003</small>; Euglenina <small>Buetschli 1884</small>; Euglenomorphales <small>Leedale 1967</small>; Colaciales <small>Smith 1938</small>]
- Family Euglenamorphidae Hollande, 1952 stat. n. Cavalier-Smith 2016 [Euglenomorphaceae; Hegneriaceae <small>Brumpt & Lavier 1924</small>]
- Family Phacidae [Phacaceae <small>Kim et al. 2010</small>]
- Family Euglenidae Bütschli 1884 [Euglenaceae <small>Dujardin 1841 emend. Kim et al. 2010</small>; Colaciaceae Smith 1933] (Mucilaginous clade)
Kostygov et al. (2021)
Phylum Euglenozoa Cavalier-Smith 1981 emend. Simpson 1997[9]
- Class Kinetoplastea Honigberg, 1963 emend. Vickerman, 1976
- Subclass Prokinetoplastia Vickerman, 2004
- Order Prokinetoplastida Vickerman, 2004
- Family Ichthyobodonidae Isaksen et al., 2007
- Family Perkinselidae Kostygov, 2021
- Subclass Metakinetoplastia Vickerman, 2004
- Order Eubodonida Vickerman 2004
- Family Bodonidae Bütschli, 1883
- Order Neobodonida Vickerman, 2004
- Family Allobodonidae Goodwin et al., 2018
- Family Neobodonidae Cavalier-Smith, 2016
- Family Rhynchomonadinae Cavalier-Smith, 2016
- Order Parabodonida Vickerman, 2004
- Family Cryptobiidae Poche, 1911 emend. Kostygov, 2021
- Family Trypanoplasmatidae Hartmann and Chagas, 1910 emend. Kostygov, 2021
- Order Trypanosomatida Kent, 1880
- Class Diplonemea Cavalier-Smith, 1993
- Order Diplonemida Cavalier-Smith, 1993
- Family Diplonemidae Cavalier-Smith, 1993
- Family Hemistasiidae Cavalier-Smith, 2016
- Family Eupelagonemidae Okamoto and Keeling, 2019
- Class Euglenida Bütschli, 1884 emend. Simpson, 1997
- Clade Olkaspira Lax and Simpson, 2020
- Clade Spirocuta Cavalier-Smith, 2016
- Clade Euglenophyceae Schoenichen, 1925 emend. Marin and Melkonian, 2003
- Order Euglenales Leedale, 1967 emend. Marin and Melkonian, 2003
- Family Euglenaceae Dujardin, 1841 emend. Kim et al., 2010 [Euglenidae <small>Dujardin, 1841</small>]
- Family Phacaceae Kim, Triemer and Shin 2010 [Phacidae <small>Kim, Triemer and Shin 2010</small>]
- Order Eutreptiales Leedale, 1967 emend. Marin and Melkonian, 2003
- Family Eutreptiaceae Hollande, 1942 [Eutreptiidae <small>Hollande, 1942</small>]
- Order Rapazida Cavalier-Smith, 2016
- Clade Anisonemia Cavalier-Smith, 2016
- Order Anisonemida Cavalier-Smith, 2016
- Clade Aphagea Cavalier-Smith, 1993 emend. Busse and Preisfeld, 2002
- Order Peranemida Cavalier-Smith, 1993
- Clade Alistosa Lax et al., 2020
- Order Petalomonadida Cavalier-Smith, 1993
- Class Symbiontida Yubuki, Edgcomb, Bernhard and Leander, 2009
External links
Notes and References
- Web site: Euglenozoa . Encyclopedia of Life . National Museum of Natural History - Smithsonian Institution . 16 January 2020.
- Patterson DJ . The Diversity of Eukaryotes . The American Naturalist . 154 . S4 . S96–S124 . October 1999 . 10527921 . 10.1086/303287 . 4367158 .
- Dorrell RG, Smith AG . Do red and green make brown?: perspectives on plastid acquisitions within chromalveolates . Eukaryotic Cell . 10 . 7 . 856–868 . July 2011 . 21622904 . 3147421 . 10.1128/EC.00326-10 .
- Simpson AG . The Identity and Composition of Euglenozoa . Archiv für Protistenkunde . 1997 . 148 . 3 . 318–328 . 10.1016/s0003-9365(97)80012-7.
- Baldauf SL, Roger AJ, Wenk-Siefert I, Doolittle WF . A kingdom-level phylogeny of eukaryotes based on combined protein data . Science . 290 . 5493 . 972–977 . November 2000 . 11062127 . 10.1126/science.290.5493.972 . 2000Sci...290..972B . Ford Doolittle .
- Simpson AG . Cytoskeletal organization, phylogenetic affinities and systematics in the contentious taxon Excavata (Eukaryota) . International Journal of Systematic and Evolutionary Microbiology . 53 . Pt 6 . 1759–1777 . November 2003 . 14657103 . 10.1099/ijs.0.02578-0 . free .
- Cavalier-Smith T . Kingdoms Protozoa and Chromista and the eozoan root of the eukaryotic tree . Biology Letters . 6 . 3 . 342–345 . June 2010 . 20031978 . 2880060 . 10.1098/rsbl.2009.0948 .
- Cavalier-Smith T . Higher classification and phylogeny of Euglenozoa . European Journal of Protistology . 56 . 250–276 . October 2016 . 27889663 . 10.1016/j.ejop.2016.09.003 . free .
- Kostygov AY, Karnkowska A, Votýpka J, Tashyreva D, Maciszewski K, Yurchenko V, Lukeš J . Euglenozoa: taxonomy, diversity and ecology, symbioses and viruses . Open Biology . 11 . 3 . 200407 . March 2021 . 33715388 . 8061765 . 10.1098/rsob.200407 .
- Cavalier-Smith T . Euglenoid pellicle morphogenesis and evolution in light of comparative ultrastructure and trypanosomatid biology: Semi-conservative microtubule/strip duplication, strip shaping and transformation . European Journal of Protistology . 61 . Pt A . 137–179 . October 2017 . 29073503 . 10.1016/j.ejop.2017.09.002 . free .