Euglenozoa Temporal range: | |
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Two Euglena | |
Scientific classification | |
Domain: | Eukaryota |
Clade: | Discoba |
Superphylum: | Discicristata |
Phylum: | Euglenozoa Cavalier-Smith, 1981 [2] |
Classes | |
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 15–40 μm (0.00059–0.00157 in) in size, although some euglenids get up to 500 μm (0.020 in) long. [3]
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. [4]
Some other euglenozoa feed through absorption, and many euglenids possess chloroplasts, the only eukaryotes outside Diaphoretickes to do so without performing kleptoplasty, [5] [ failed verification ] 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. [4]
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 . [6]
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. [7]
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. [8] Both probably belong to a larger group of eukaryotes called the Excavata. [9] This grouping, though, has been challenged. [10]
The phylogeny based on the work of Cavalier-Smith (2016): [11]
Euglenozoa phylogeny – Cavalier-Smith (2016) | |||||||||||||||||||||||||||||||||||||||||||||||
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A consensus phylogeny following the review by Kostygov et al. (2021): [7]
Euglenozoa phylogeny – Kostygov et al. (2021) | |||||||||
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The following classification of Euglenozoa is as described by Cavalier-Smith in 2016, [11] modified to include the new subphylum Plicomonada according to Cavalier-Smith et al (2017). [12]
Phylum EuglenozoaCavalier-Smith 1981 emend. Simpson 1997 [Euglenobionta]
Phylum EuglenozoaCavalier-Smith 1981 emend. Simpson 1997 [7]
The alveolates are a group of protists, considered a major clade and superphylum within Eukarya. They are currently grouped with the stramenopiles and Rhizaria among the protists with tubulocristate mitochondria into the SAR supergroup.
Euglenids are one of the best-known groups of flagellates, which are excavate eukaryotes of the phylum Euglenophyta and their cell structure is typical of that group. They are commonly found in freshwater, especially when it is rich in organic materials, with a few marine and endosymbiotic members. Many euglenids feed by phagocytosis, or strictly by diffusion. A monophyletic group consisting of the mixotrophic Rapaza viridis and the two groups Eutreptiales and Euglenales have chloroplasts and produce their own food through photosynthesis. This group is known to contain the carbohydrate paramylon.
Kinetoplastida is a group of flagellated protists belonging to the phylum Euglenozoa, and characterised by the presence of a distinctive organelle called the kinetoplast, a granule containing a large mass of DNA. The group includes a number of parasites responsible for serious diseases in humans and other animals, as well as various forms found in soil and aquatic environments. The organisms are commonly referred to as "kinetoplastids" or "kinetoplasts".
Amoebozoa is a major taxonomic group containing about 2,400 described species of amoeboid protists, often possessing blunt, fingerlike, lobose pseudopods and tubular mitochondrial cristae. In traditional classification schemes, Amoebozoa is usually ranked as a phylum within either the kingdom Protista or the kingdom Protozoa. In the classification favored by the International Society of Protistologists, it is retained as an unranked "supergroup" within Eukaryota. Molecular genetic analysis supports Amoebozoa as a monophyletic clade. Modern studies of eukaryotic phylogenetic trees identify it as the sister group to Opisthokonta, another major clade which contains both fungi and animals as well as several other clades comprising some 300 species of unicellular eukaryotes. Amoebozoa and Opisthokonta are sometimes grouped together in a high-level taxon, variously named Unikonta, Amorphea or Opimoda.
Phaeodarea or Phaeodaria is a group of amoeboid cercozoan organisms. They are traditionally considered radiolarians, but in molecular trees do not appear to be close relatives of the other groups, and are instead placed among the Cercozoa. They are distinguished by the structure of their central capsule and by the presence of a phaeodium, an aggregate of waste particles within the cell.
The metamonads are a large group of flagellate amitochondriate microscopic eukaryotes. Their composition is not entirely settled, but they include the retortamonads, diplomonads, and possibly the parabasalids and oxymonads as well. These four groups are all anaerobic, occurring mostly as symbiotes or parasites of animals, as is the case with Giardia lamblia which causes diarrhea in mammals.
Streptophyta, informally the streptophytes, is a clade of plants. The composition of the clade varies considerably between authors, but the definition employed here includes land plants and all green algae except the Chlorophyta and the more basal Prasinodermophyta.
Yellow-green algae or the Xanthophyceae (xanthophytes) are an important group of heterokont algae. Most live in fresh water, but some are found in marine and soil habitats. They vary from single-celled flagellates to simple colonial and filamentous forms. Xanthophyte chloroplasts contain the photosynthetic pigments chlorophyll a, chlorophyll c, β-carotene, and the carotenoid diadinoxanthin. Unlike other Stramenopiles (heterokonts), their chloroplasts do not contain fucoxanthin, which accounts for their lighter colour. Their storage polysaccharide is chrysolaminarin. Xanthophyte cell walls are produced of cellulose and hemicellulose. They appear to be the closest relatives of the brown algae.
The Ichthyosporea are a small group of Opisthokonta in Eukaryota, mostly parasites of fish and other animals.
The Archamoebae are a group of protists originally thought to have evolved before the acquisition of mitochondria by eukaryotes. They include genera that are internal parasites or commensals of animals. A few species are human pathogens, causing diseases such as amoebic dysentery. The other genera of archamoebae live in freshwater habitats and are unusual among amoebae in possessing flagella. Most have a single nucleus and flagellum, but the giant amoeba Pelomyxa has many of each.
Monadofilosa is a grouping of Cercozoa. These organisms are single-celled amoeboid protists.
Jakobids are an order of free-living, heterotrophic, flagellar eukaryotes in the supergroup Excavata. They are small, and can be found in aerobic and anaerobic environments. The order Jakobida, believed to be monophyletic, consists of only twenty species at present, and was classified as a group in 1993. There is ongoing research into the mitochondrial genomes of jakobids, which are unusually large and bacteria-like, evidence that jakobids may be important to the evolutionary history of eukaryotes.
Dictyochophyceae sensu lato is a photosynthetic lineage of heterokont algae.
The sarcomonads or class Sarcomonadea are a group of amoeboid biciliate protists in the phylum Cercozoa. They are characterized by a propensity to move through gliding on their posterior cilium or through filopodia, a lack of scales or external theca, a soft cell surface without obvious cortical filamentous or membranous skeleton, two cilia without scales or hairs, tubular mitochondrial cristae, near-spherical extrusomes, and a microbody attached to the nucleus.
Bodonida is an order of kinetoplastid flagellate excavates. It contains the genera Bodo and Rhynchomonas, relatives to the parasitic trypanosomes. This order also contains the colonial genus Cephalothamnium.
A protist is any eukaryotic organism 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; 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". In the 21st century, the classification shifted toward a two-kingdom system of protists: Chromista and Protozoa.
Postgaardia is a proposed basal clade of flagellate Euglenozoa, following Thomas Cavalier-Smith. As of April 2023, the Interim Register of Marine and Nonmarine Genera treats the group as a subphylum. A 2021 review of Euglenozoa places Cavalier-Smith's proposed members of Postgaardia in the class Symbiontida. As Euglenozoans may be basal eukaryotes, the Postgaardia may be key to studying the evolution of Eukaryotes, including the incorporation of eukaryotic traits such as the incorporation of alphaproteobacterial mitochondrial endosymbionts.
Ventrifilosa is a highly diverse group of phagotrophic protists that glide through their flagella and emit filose pseudopods from their ventral side for feeding. Because of their mixture of amoeba and flagellate characteristics, they are amoeboflagellates. Members of this group are the Imbricatea, Sarcomonadea and Thecofilosea.
Gyrista is a phylum of heterokont protists containing three diverse groups: the mostly photosynthetic Ochrophyta, the parasitic Pseudofungi, and the recently described group of nanoflagellates known as Bigyromonada. Members of this phylum are characterized by the presence of a helix or a double helix/ring system in the ciliary transition region.
The glissomonads are a group of bacterivorous gliding flagellated protists that compose the order Glissomonadida, in the amoeboflagellate phylum Cercozoa. They comprise a vast, largely undescribed diversity of soil and freshwater organisms. They are the sister group to cercomonads; the two orders form a solid clade of gliding soil-dwelling flagellates called Pediglissa.