| Percolozoa | |
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| The three different stages of N. fowleri | |
Scientific classification  | |
| Domain: | Eukaryota |
| Clade: | Diphoda |
| Clade: | Discoba |
| Superphylum: | Discicristata |
| Phylum: | Percolozoa Cavalier-Smith 1991 |
| Classes | |
| Synonyms | |
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The Percolozoa are a group of colourless, non-photosynthetic Excavata, including many that can transform between amoeboid, flagellate, and cyst stages.
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Most Percolozoa are found as bacterivores in soil, fresh water and occasionally in the ocean. The only member of this group that is infectious to humans is Naegleria fowleri , the causative agent of the often fatal disease amoebic meningitis. The group is closely related to the Euglenozoa, and share with them the unusual characteristic of having mitochondria with discoid cristae. The presence of a ventral feeding groove in the flagellate stage, as well as other features, suggests that they are part of the Excavata group.
The amoeboid stage is roughly cylindrical, typically around 20–40 μm in length. They are traditionally considered lobose amoebae, but are not related to the others, and unlike them, do not form true lobose pseudopods. Instead, they advance by eruptive waves, where hemispherical bulges appear from the front margin of the cell, which is clear. The flagellate stage is slightly smaller, with two or four anterior flagella anterior to the feeding groove.
Usually, the amoeboid form is taken when food is plentiful, and the flagellate form is used for rapid locomotion. However, not all members are able to assume both forms. The genera Percolomonas , Lyromonas , and Psalteriomonas are known only as flagellates, while Vahlkampfia , Pseudovahlkampfia , and most acrasids do not have flagellate stages. As mentioned above, under unfavourable conditions, the acrasids aggregate to form sporangia. These are superficially similar to the sporangia of the dictyostelids, but the amoebae only aggregate as individuals or in small groups and do not die to form the stalk.
These are collectively referred to as schizopyrenids, amoeboflagellates, or vahlkampfids. They also include the acrasids, a group of social amoebae that aggregate to form sporangia. The entire group is usually called the Heterolobosea, but this may be restricted to members with amoeboid stages.
One Heterolobosea classification system is: [1]
Pleurostomum flabellatum has recently been added to Heterolobosea. [2]
Based on the cladogram from Tolweb and updated by Pánek and Čepička 2014. [3] [4]
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Phylum Percolozoa Cavalier-Smith 1991 [6]
The Heterolobosea were first defined by Page and Blanton in 1985 [7] as a class of amoebae, and so only included those forms with amoeboid stages. Cavalier-Smith created the phylum Percolozoa for the extended group, together with the enigmatic flagellate Stephanopogon . [8]
Cavalier-Smith maintained the Heterolobosea as a class for amoeboid forms. He has defined Percolozoa as "Heterolobosea plus Percolatea classis nov." [9]
A flagellate is a cell or organism with one or more whip-like appendages called flagella. The word flagellate also describes a particular construction characteristic of many prokaryotes and eukaryotes and their means of motion. The term presently does not imply any specific relationship or classification of the organisms that possess flagella. However, several derivations of the term "flagellate" are more formally characterized.
Excavata is an extensive and diverse but paraphyletic group of unicellular Eukaryota. The group was first suggested by Simpson and Patterson in 1999 and the name latinized and assigned a rank by Thomas Cavalier-Smith in 2002. It contains a variety of free-living and symbiotic protists, and includes some important parasites of humans such as Giardia and Trichomonas. Excavates were formerly considered to be included in the now obsolete Protista kingdom. They were distinguished from other lineages based on electron-microscopic information about how the cells are arranged. They are considered to be a basal flagellate lineage.
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, named Amorphea. Amoebozoa includes many of the best-known amoeboid organisms, such as Chaos, Entamoeba, Pelomyxa and the genus Amoeba itself. Species of Amoebozoa may be either shelled (testate) or naked, and cells may possess flagella. Free-living species are common in both salt and freshwater as well as soil, moss and leaf litter. Some live as parasites or symbionts of other organisms, and some are known to cause disease in humans and other organisms.
The metamonads are a large group of flagellate amitochondriate microscopic eukaryotes. They include the retortamonads, diplomonads, parabasalids, oxymonads, and a range of more poorly studied taxa, most of which are free-living flagellates. All metamonads are anaerobic, and most members of the four groups listed above are symbiotes or parasites of animals, as is the case with Giardia lamblia which causes diarrhea in mammals.
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.
The Oxymonads are a group of flagellated protists found exclusively in the intestines of animals, mostly termites and other wood-eating insects. Along with the similar parabasalid flagellates, they harbor the symbiotic bacteria that are responsible for breaking down cellulose. There is no evidence for presence of mitochondria in oxymonads and three species have been shown to completely lack any molecular markers of mitochondria.
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.
The family Acrasidae is a family of slime molds which belongs to the excavate group Percolozoa. The name element acrasio- comes from the Greek akrasia, meaning "acting against one's judgement". This group consists of cellular slime molds.
Cafeteria is a genus of marine bicosoecid described in 1988 by Tom Fenchel and D. J. Patterson. It was created after the discovery of a new species, Cafeteria roenbergensis, a tiny (5–10 μm) eukaryotic organism that is eaten by protozoa and small invertebrates. The name is meant to indicate the importance of the genus in the food web.
Anaeromonadea, also known as Preaxostyla, is a class of excavate protists, comprising the oxymonads, Trimastix, and Paratrimastix. This group is studied as a model system for reductive evolution of mitochondria, because it includes both organisms with anaerobic mitochondrion-like organelles, and those that have completely lost their mitochondria.
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.
Psalteriomonas is a genus of excavates in the group of Heterolobosea. The genus was first discovered and named in 1990. It contains amoeboflagellate cells that live in freshwater anaerobic sediments all over the world. The microtubule-organizing ribbon and the associated microfibrillar bundles of the mastigote system is the predominant feature in Psalteriomonas. This harp-shaped complex gives rise to the name of this genus. Psalteriomonasforms an endosymbiotic relationship with methanogenic bacteria, especially with Methanobacterium formicicum There are currently three species in this genus: P. lanterna, P. vulgaris, and P. magna.
The vampyrellids, colloquially known as vampire amoebae, are a group of free-living predatory amoebae classified as part of the lineage Endomyxa. They are distinguished from other groups of amoebae by their irregular cell shape with propensity to fuse and split like plasmodial organisms, and their life cycle with a digestive cyst stage that digests the gathered food. They appear worldwide in marine, brackish, freshwater and soil habitats. They are important predators of an enormous variety of microscopic organisms, from algae to fungi and animals. They are also known as aconchulinid amoebae.
An amoeba, often called an amoeboid, is a type of cell or unicellular organism with the ability to alter its shape, primarily by extending and retracting pseudopods. Amoebae do not form a single taxonomic group; instead, they are found in every major lineage of eukaryotic organisms. Amoeboid cells occur not only among the protozoa, but also in fungi, algae, and animals.
Stygiella /ˌstɪ.d͡ʒiˈɛ.lə/ is a genus of free-living marine flagellates belonging to the family Stygiellidae in the jakobids (excavata).
Anaeramoeba is a genus of anaerobic protists of uncertain phylogenetic position, first described in 2016.
Stygiellidae is a family of free-living marine flagellates belonging to the order Jakobida, a deep-branching lineage within the eukaryotic supergroup Discoba. They are unicellular organisms that commonly inhabit anoxic, sulfide-rich and ammonium-rich marine habitats worldwide.
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.