Amorphea | |
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Scientific classification | |
Domain: | Eukaryota |
Clade: | Amorphea Adl et al., 2012 [1] |
Subgroups | |
Synonyms | |
Amorphea [1] is a taxonomic supergroup that includes the basal Amoebozoa and Obazoa. That latter contains the Opisthokonta, which includes the Fungi, Animals and the Choanomonada, or Choanoflagellates. The taxonomic affinities of the members of this clade were originally described and proposed by Thomas Cavalier-Smith in 2002. [2] [4]
The International Society of Protistologists, the recognised body for taxonomy of protozoa, recommended in 2012 that the term Unikont be changed to Amorphea because the name "Unikont" is based on a hypothesized synapomorphy that the ISOP authors and other scientists later rejected. [1] [5]
It includes amoebozoa, opisthokonts, [6] [7] and Apusomonada. [8]
Thomas Cavalier-Smith proposed two new phyla: Sulcozoa, which consists of the subphyla Apusozoa (Apusomonadida and Breviatea), and Varisulca, which includes the subphyla Diphyllatea, Discocelida, Mantamonadidae, Planomonadida and Rigifilida. [9]
Further work by Cavalier-Smith showed that Sulcozoa is paraphyletic. [10] Apusozoa also appears to be paraphyletic. Varisulca has been redefined to include planomonads, Mantamonas and Collodictyon. A new taxon has been created - Glissodiscea - for the planomonads and Mantamonas. Again, the validity of this revised taxonomy awaits confirmation.
Amoebozoa seems to be monophyletic with two major branches: Conosa and Lobosa. Conosa is divided into the aerobic infraphylum Semiconosia (Mycetozoa and Variosea) and secondarily anaerobic Archamoebae. Lobosa consists entirely of non-flagellated lobose amoebae and has been divided into two classes: Discosea, which have flattened cells, and Tubulinea, which has predominantly tube-shaped pseudopodia. [11]
The group includes eukaryotic cells that, for the most part, have a single emergent flagellum, or are amoebae with no flagella. The unikonts include opisthokonts (animals, fungi, and related forms) and Amoebozoa. By contrast, other well-known eukaryotic groups, which more often have two emergent flagella (although there are many exceptions), are often referred to as bikonts. Bikonts include Archaeplastida (plants and relatives) and SAR supergroup, the Cryptista, Haptista, Telonemia and picozoa.
Eukaryotes |
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2200 mya |
One view of the great kingdoms and their stem groups. [12] [13] [14] [15] The Metamonada are hard to place, being sister possibly to Discoba, possibly to Malawimonada. [15]
The unikonts have a triple-gene fusion that is lacking in the bikonts. The three genes that are fused together in the unikonts, but not bacteria or bikonts, encode enzymes for synthesis of the pyrimidine nucleotides: carbamoyl phosphate synthase, dihydroorotase, aspartate carbamoyltransferase. This must have involved a double fusion, a rare pair of events, supporting the shared ancestry of Opisthokonta and Amoebozoa.
Cavalier-Smith [2] originally proposed that unikonts ancestrally had a single flagellum and single basal body. This is unlikely, however, as flagellated opisthokonts, as well as some flagellated Amoebozoa, including Breviata , actually have two basal bodies, as in typical 'bikonts' (even though only one is flagellated in most unikonts). This paired arrangement can also be seen in the organization of centrioles in typical animal cells. In spite of the name of the group, the common ancestor of all 'unikonts' was probably a cell with two basal bodies.
Cercozoa is a phylum of diverse single-celled eukaryotes. They lack shared morphological characteristics at the microscopic level, and are instead united by molecular phylogenies of rRNA and actin or polyubiquitin. They were the first major eukaryotic group to be recognized mainly through molecular phylogenies. They are the natural predators of many species of bacteria. They are closely related to the phylum Retaria, comprising amoeboids that usually have complex shells, and together form a supergroup called Rhizaria.
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.
The opisthokonts are a broad group of eukaryotes, including both the animal and fungus kingdoms. The opisthokonts, previously called the "Fungi/Metazoa group", are generally recognized as a clade. Opisthokonts together with Apusomonadida and Breviata comprise the larger clade Obazoa.
A bikont is any of the eukaryotic organisms classified in the group Bikonta. Many single-celled and multi-celled organisms are members of the group, and these, as well as the presumed ancestor, have two flagella.
The Apusozoa are a paraphyletic phylum of flagellate eukaryotes. They are usually around 5–20 μm in size, and occur in soils and aquatic habitats, where they feed on bacteria. They are grouped together based on the presence of an organic shell or theca under the dorsal surface of the cell.
Discosea is a class of Amoebozoa, consisting of naked amoebae with a flattened, discoid body shape. Members of the group do not produce tubular or subcylindrical pseudopodia, like amoebae of the class Tubulinea. When a discosean is in motion, a transparent layer called hyaloplasm forms at the leading edge of the cell. In some discoseans, short "subpseudopodia" may be extended from this hyaloplasm, but the granular contents of the cell do not flow into these, as in true pseudopodia. Discosean amoebae lack hard shells, but some, like Cochliopodium and Korotnevella secrete intricate organic scales which may cover the upper (dorsal) surface of the cell. No species have flagella or flagellated stages of life.
Telonemia is a phylum of microscopic eukaryotes commonly known as telonemids. They are unicellular free-living flagellates with a unique combination of cell structures, including a highly complex cytoskeleton unseen in other eukaryotes.
Corticata, in the classification of eukaryotes, is a clade suggested by Thomas Cavalier-Smith to encompass the eukaryote supergroups of the following two groups:
Breviata anathema is a single-celled flagellate amoeboid eukaryote, previously studied under the name Mastigamoeba invertens. The cell lacks mitochondria, much like the pelobionts to which the species was previously assigned, but has remnant mitochondrial genes, and possesses an organelle believed to be a modified anaerobic mitochondrion, similar to the mitosomes and hydrogenosomes found in other eukaryotes that live in low-oxygen environments.
Conosa is a grouping of Amoebozoa. It is subdivided into three groups: Archamoeba, Variosea and Mycetozoa.
Ancyromonadida or Planomonadida is a small group of biflagellated protists found in the soil and in aquatic habitats, where they feed on bacteria. Includes freshwater or marine organisms, benthic, dorsoventrally compressed and with two unequal flagellae, each emerging from a separate pocket. The apical anterior flagellum can be very thin or end in the cell membrane, while the posterior flagellum is long and is inserted ventrally or laterally. The cell membrane is supported by a thin single-layered theca and the mitochondrial crests are discoidal/flat.
Diaphoretickes is a major group of eukaryotic organisms, with over 400,000 species. The majority of the earth's biomass that carries out photosynthesis belongs to Diaphoretickes.
Breviatea, commonly known as breviate amoebae, are a group of free-living, amitochondriate protists with uncertain phylogenetic position. They are biflagellate, and can live in anaerobic (oxygen-free) environments. They are currently placed in the Obazoa clade. They likely do not possess vinculin proteins. Their metabolism relies on fermentative production of ATP as an adaptation to their low-oxygen environment.
Varisulca was a proposed basal Podiate taxon. It encompassed several lineages of heterotrophic protists, most notably the ancyromonads (planomonads), collodictyonids (diphylleids), rigifilids and mantamonadids. Recent evidence suggests that the latter three are closely related to each other, forming a clade called CRuMs, but that this is unlikely to be specifically related to ancyromonads
Obazoa is a proposed sister clade of Amoebozoa. The term Obazoa is based on the OBA acronym for Opisthokonta, Breviatea, and Apusomonadida, the group's three constituent clades.
The Orthokaryotes are a proposed Eukaryote clade consisting of the Jakobea and the Neokaryotes. Together with its sister Discicristata it forms a basal Eukaryote clade. They are characterized by stacked Golgi, orthogonal centrioles, and two opposite posterior ciliary roots.
The neokaryotes are a proposed eukaryote clade consisting of the unikonts and the bikonts as sister of for instance the Jakobea. It arises because the Euglenozoa, Percolozoa, Tsukubea, and Jakobea are seen in this view as more basal eukaryotes. These four groups, are traditionally grouped together in the Discoba. However, the Discoba may well be paraphyletic as the neokaryotes may have emerged in them.
The Scotokaryotes (Cavalier-Smith) is a proposed basal Neokaryote clade as sister of the Diaphoretickes. Basal Scotokaryote groupings are the Metamonads, the Malawimonas and the Podiata. In this phylogeny the Discoba are sometimes seen as paraphyletic and basal Eukaryotes.
Mantamonads are a group of free-living heterotrophic flagellates that move primarily by gliding on surfaces. They are classified as one genus Mantamonas in the monotypic family Mantamonadidae, order Mantamonadida and class Glissodiscea. Previously, they were classified in Apusozoa as sister of the Apusmonadida on the basis of rRNA analyses. However, mantamonads are currently placed in CRuMs on the basis of phylogenomic analyses that identify their closest relatives as the Diphylleida and Rigifilida.
An amoeboflagellate is any eukaryotic organism capable of behaving as an amoeba and as a flagellate at some point during their life cycle. Amoeboflagellates present both pseudopodia and at least one flagellum, often simultaneously.