Evosea | |
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Dictyostelium discoideum | |
Scientific classification | |
Domain: | Eukaryota |
Phylum: | Amoebozoa |
Clade: | Evosea Kang et al. 2017 [1] |
Clades [1] [2] | |
Evosea is a diverse clade of amoeboid protists discovered through molecular analyses. Along with Tubulinea and Discosea, Evosea is one of the three major groups within Amoebozoa, an important clade of eukaryotic organisms. It contains unicellular organisms that display a wide variety of life cycles and cell shapes, including amoebae, flagellates and different kinds of slime molds.
Evosea is a strongly supported clade of eukaryotes containing four large groups of amoebozoans: Eumycetozoa or "true" slime molds, [3] Variosea, Cutosea and Archamoebae. It is defined on a node-based approach as the least-inclusive clade containing Dictyostelium discoideum (a true slime mold), Protostelium nocturnum (a variosean), Squamamoeba japonica (a cutosean), and Entamoeba histolytica (an archamoeba). [1]
Within Evosea, organisms can vary across almost the entire range of morphologies seen in Amoebozoa. Many members have complex life cycles that include amoebae, flagellates and fruiting stages. Some species appear to be exclusively flagellates, with no amoeboid features. [2]
Evosea is a clade discovered in 2017 through a phylogenomic study by Senghuo Kang and coauthors, published in the journal Molecular Biology and Evolution . [1] Since its discovery, it has been supported by independent analyses. [4] [5] [6] As of 2019, it is accepted by the International Society of Protistologists as part of the modern cladistic classification of eukaryotes. [2] The name 'Evosea' is partly an acronym of the major members of the clade: Eumycetozoa (E), Variosea (v), Squamamoebidae and Sapocribridae (s), and Archamoebae (a). [1]
Evosea is composed of two sister clades: Cutosea, a small group of solitary amoebae, and Conosa, a larger group that contains the archamoebae, true slime molds and Variosea. Evosea, along with Tubulinea and Discosea, compose the entirety of Amoebozoa. The branching order of these three basal groups is still unresolved: either Evosea groups with Tubulinea (in a clade known as Tevosa), [1] or with Discosea (in a clade called Divosa). The following cladogram is based on a 2022 analysis, which resulted in the Divosa hypothesis: [6]
Amorphea |
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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.
Pelomyxa is a genus of giant flagellar amoebae, usually 500–800 μm but occasionally up to 5 mm in length, found in anaerobic or microaerobic bottom sediments of stagnant freshwater ponds or slow-moving streams.
Lobosa is a taxonomic group of amoebae in the phylum Amoebozoa. Most lobosans possess broad, bluntly rounded pseudopods, although one genus in the group, the recently discovered Sapocribrum, has slender and threadlike (filose) pseudopodia. In current classification schemes, Lobosa is a subphylum, composed mainly of amoebae that have lobose pseudopods but lack cilia or flagella.
Amorphea 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.
The Tubulinea are a major grouping of Amoebozoa, including most of the more familiar amoebae genera like Amoeba, Arcella, Difflugia and Hartmannella.
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.
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.
Eumycetozoa, or true slime molds, is a diverse group of protists that behave as slime molds and develop fruiting bodies, either as sorocarps or as sporocarps. It is a monophyletic group or clade within the phylum Amoebozoa that contains the myxogastrids, dictyostelids and protosporangiids.
Conosa is a grouping of Amoebozoa. It is subdivided into three groups: Archamoeba, Variosea and Mycetozoa.
Thecofilosea is a class of unicellular testate amoebae belonging to the phylum Cercozoa. They are amoeboflagellates, organisms with flagella and pseudopodia, distinguished from other cercozoa by their scale-lacking test composed of organic material. They are closely related to the Imbricatea, a group of testate amoebae with tests composed of inorganic silica scales.
Holozoa is a clade of organisms that includes animals and their closest single-celled relatives, but excludes fungi and all other organisms. Together they amount to more than 1.5 million species of purely heterotrophic organisms, including around 300 unicellular species. It consists of various subgroups, namely Metazoa and the protists Choanoflagellata, Filasterea, Pluriformea and Ichthyosporea. Along with fungi and some other groups, Holozoa is part of the Opisthokonta, a supergroup of eukaryotes. Choanofila was previously used as the name for a group similar in composition to Holozoa, but its usage is discouraged now because it excludes animals and is therefore paraphyletic.
Trichosphaerium is a genus of amoebozoan protists that present extraordinary morphological transformations, both in size and shape, during their life cycle. They can present a test that may or may not be covered in spicules. They are related to the family Microcoryciidae, which contains other amoebae with tests, within the clade Corycidia of the phylum Amoebozoa.
The apusomonads are a group of protozoan zooflagellates that glide on surfaces, and mostly consume prokaryotes. They are of particular evolutionary interest because they appear to be the sister group to the Opisthokonts, the clade that includes both animals and fungi. Together with the Breviatea, these form the Obazoa clade.
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.
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.
The Flabellinia are a subclass of Amoebozoa. During locomotion the cells are flattened and have a clear layer called hyaloplasm along the front margin. Some form slender subpseudopodia projecting outward from the hyaloplasm, but the cell mass does not flow into these as in true pseudopodia, and advances without a definite central axis as in the Tubulinea. They also lack distinctive features like shells and flagella, and are united mainly by evidence from molecular trees.
Cutosea is a small group of marine amoeboid protists proposed in 2016. It is a monotypic class of Amoebozoa containing the order Squamocutida. Cutosean organisms are characterized by a cell coat of microscales separated from the cell membrane. Three genera, Armaparvus, Sapocribrum and Squamamoeba, belong to this group, distributed in two families.
Sainouroidea is a group of microscopic protists belonging to the supergroup Rhizaria, both discovered through molecular phylogenetic analyses. It contains amoeboid flagellates with two flagella. They are either free-living, mostly on fecal matter, or live inside the gut of animals. Among these amoebae, one lineage has independently evolved aggregative multicellularity similarly to slime moulds.
Syssomonas is a monotypic genus of unicellular flagellated protists containing the species Syssomonas multiformis. It is a member of Pluriformea inside the lineage of Holozoa, a clade containing animals and their closest protistan relatives. It lives in freshwater habitats. It has a complex life cycle that includes unicellular amoeboid and flagellated phases, as well as multicellular aggregates, depending on the growth medium and nutritional state.
Corycidia is a clade of amoeboid protists within the eukaryotic supergroup Amoebozoa. It contains all amoebae of the families Microcoryciidae, which was previously regarded as Arcellinida, and Trichosphaeriidae, which contains the sole genus Trichosphaerium.