| Percolozoa | |
|---|---|
 | |
| The three different stages of N. fowleri | |
Scientific classification  | |
| Domain: | Eukaryota |
| Clade: | Diphoda |
| Clade: | Discoba |
| Superphylum: | Discicristata |
| Phylum: | Percolozoa Cavalier-Smith 1991 |
| Classes | |
| |
| Synonyms | |
| |
The Percolozoa are a group of colourless, non-photosynthetic Excavata, including many that can transform between amoeboid, flagellate, and cyst stages.
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]
| |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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, the term "flagellate" is included in other terms which 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, variously named Unikonta, Amorphea or Opimoda.
The Rhizaria are a diverse and species-rich supergroup of mostly unicellular eukaryotes. Except for the Chlorarachniophytes and three species in the genus Paulinella in the phylum Cercozoa, they are all non-photosynthethic, but many foraminifera and radiolaria have a symbiotic relationship with unicellular algae. A multicellular form, Guttulinopsis vulgaris, a cellular slime mold, has been described. This group was used by Cavalier-Smith in 2002, although the term "Rhizaria" had been long used for clades within the currently recognized taxon. Being described mainly from rDNA sequences, they vary considerably in form, having no clear morphological distinctive characters (synapomorphies), but for the most part they are amoeboids with filose, reticulose, or microtubule-supported pseudopods. In the absence of an apomorphy, the group is ill-defined, and its composition has been very fluid. Some Rhizaria possess mineral exoskeletons, which are in different clades within Rhizaria made out of opal, celestite, or calcite. Certain species can attain sizes of more than a centimeter with some species being able to form cylindrical colonies approximately 1 cm in diameter and greater than 1 m in length. They feed by capturing and engulfing prey with the extensions of their pseudopodia; forms that are symbiotic with unicellular algae contribute significantly to the total primary production of the ocean.
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.
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 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.
Monadofilosa is a grouping of Cercozoa. These organisms are single-celled amoeboid protists.
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 is a class of excavates, comprising the oxymonads and Trimastix.
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 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.
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.