Testate amoebae (formerly thecamoebians, Testacea or Thecamoeba) are a polyphyletic group of unicellular amoeboid protists, which differ from naked amoebae in the presence of a test that partially encloses the cell, with an aperture from which the pseudopodia emerge, that provides the amoeba with shelter from predators and environmental conditions.
The test of some species is produced entirely by the amoeba and may be organic, siliceous or calcareous depending on the species (autogenic tests), whereas in other cases the test is made up of particles of sediment collected by the amoeba which are then agglutinated together by secretions from within the cell (xenogenic tests). A few taxa (Hyalosphenidae) can build either type, depending on the circumstances and availability of foreign material. [1]
The assemblage referred to as "testate amoebae" is actually composed of several, unrelated groups of organisms. However, some features they all share that have been used to group them together include the presence of a test (regardless of its composition) and pseudopodia that do not anastomose. [2]
Testate amoebae can be found in most freshwater environments, including lakes, rivers, cenotes, [3] as well as mires and soils.
The strong and resistant nature of the tests allows them to be preserved long after the amoeba has died. These characteristics, along with the sensitivity that some species display to changes in environmental conditions (such as temperature, pH, and conductivity), has sparked their use as bioindicators and paleoclimate proxies in recent years. [4]
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Testate amoebae are a polyphyletic assemblage. The main testate amoebae groups are the lobose Tubulinea, which include Arcellinida, Difflugina and Phryganellina (within the Amoebozoa), [5] and the filose Euglyphida (within the SAR supergroup), [6] although there are smaller groups that also include other testate amoebae. [7]
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The following table includes a few examples of testate amoebae genera, and reflects their position within the classification by Adl et al. (2012), [7] where five supergroups (Amoebozoa, Opisthokonta, Excavata, SAR and Archaeplastida) were proposed to classify all eukaryotes. This classification purposefully avoids the use of Linnaean higher category names (phylum, class, order, family). While it has been noted that the names that Adl et al. provide for the clades may result confusing or uninformative regarding the relative degree of phenotypic distinctiveness amongst groups when used in isolation, [8] this system avoids creating superfluous ranks where unnecessary and provides stable group names that can be retained even when a group is moved to a different lineage, as is often the case with protists, as their classification remains in constant review. [7]
Amoebozoa | Tubulinea | Arcellinida | Arcellina | Amphizonella - Arcella - Microchlamys - Microcorycia - Spumochlamys | |||
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Difflugina | Bullinularia - Centropyxis - Difflugia - Distomatopyxis - Heleopera - Hyalosphenia - Lesquereusia - Nebela - Paraquadrula - Pontigulasia - Plagiopyxis - Quadrulella - Trigonopyxis | ||||||
Phryganellina | Cryptodifflugia - Phryganella - Wailesella | ||||||
Discosea | Himatismenida | Cochliopodium | |||||
SAR Supergroup | Stramenopila | Labyrinthulomycetes | Amphitremida | Amphitrema - Archerella | |||
Rhizaria | Cercozoa | Thecofilosea | Cryomonadida | Rhizaspididae | Capsellina - Rhizaspis - Rhogostoma | ||
Ventricleftida | Ventrifissura - Verrucomonas | ||||||
Imbricatea | Silicofilosea | Euglyphida | Euglyphidae | Euglypha - Scutiglypha | |||
Assulinidae | Assulina - Placocista - Valkanovia | ||||||
Trinematidae | Corythion - Playfairina - Puytoracia - Trinema | ||||||
Cyphoderidae | Campascus - Corythionella - Cyphoderia - Messemvriella - Pseudocorythion - Schaudinnula. | ||||||
Paulinellidae | Ovulinata - Paulinella |
Traditionally, those species that form large networks of anastomosing pseudopodia, despite some of them having tests, are not counted amongst testate amoebae; this comprises genus Gromia and the Foraminifera (both in Rhizaria). [2]
The Thecamoebida (Amoebozoa), with the genus Thecamoeba , despite their name, do not have tests.
Euglyphid testate amoebae are closely related to the Foraminifera. [9]
A pseudopod or pseudopodium is a temporary arm-like projection of a eukaryotic cell membrane that is developed in the direction of movement. Filled with cytoplasm, pseudopodia primarily consist of actin filaments and may also contain microtubules and intermediate filaments. Pseudopods are used for motility and ingestion. They are often found in amoebas.
Foraminifera are single-celled organisms, members of a phylum or class of amoeboid protists characterized by streaming granular ectoplasm for catching food and other uses; and commonly an external shell of diverse forms and materials. Tests of chitin are believed to be the most primitive type. Most foraminifera are marine, the majority of which live on or within the seafloor sediment, while a smaller number float in the water column at various depths, which belong to the suborder Globigerinina. Fewer are known from freshwater or brackish conditions, and some very few (nonaquatic) soil species have been identified through molecular analysis of small subunit ribosomal DNA.
The Cercozoa are a group of single-celled eukaryotes. They lack shared morphological characteristics at the microscopic level, being defined by molecular phylogenies of rRNA and actin or polyubiquitin. They are the natural predators of many species of microbacteria and Archea.
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 most classification schemes, Amoebozoa is 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. Most phylogenetic trees identify it as the sister group to Opisthokonta, another major clade which contains both fungi and animals as well as some 300 species of unicellular protists. Amoebozoa and Opisthokonta are sometimes grouped together in a high-level taxon, variously named Unikonta, Amorphea or Opimoda.
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 euglyphids are a prominent group of filose amoebae that produce shells or tests from siliceous scales, plates, and sometimes spines. These elements are created within the cell and then assembled on its surface in a more or less regular arrangement, giving the test a textured appearance. There is a single opening for the long slender pseudopods, which capture food and pull the cell across the substrate.
The tectofilosids are a group of filose amoebae with shells. These are composed of organic materials and sometimes collected debris, in contrast to the euglyphids, which produce shells from siliceous scales. The shell usually has a single opening, but in Amphitrema and a few other genera it has two on opposite ends. The cell itself occupies most of the shell. They are most often found on marsh plants such as Sphagnum.
Chaos is a genus of single-celled amoeboid organisms in the family Amoebidae. The largest and best-known species, the so-called "giant amoeba" Chaos carolinensis, can reach lengths of 5 mm, although most specimens fall between 1 and 3 mm.
The Amoebidae are a family of Amoebozoa, including naked amoebae that produce multiple pseudopodia of indeterminate length. These are roughly cylindrical with granular endoplasm and no subpseudopodia, as found in other members of the class Tubulinea. During locomotion one pseudopod typically becomes dominant and the others are retracted as the body flows into it. In some cases the cell moves by "walking", with relatively permanent pseudopodia serving as limbs.
Difflugia is the largest genus of Arcellinida, one of several groups of Tubulinea within the eukaryote supergroup Amoebozoa. Arcellinida species produce shells or tests from mineral particles or biogeonic elements and are thus commonly referred to as testate amoebae or shelled amoebae. Difflugia are particularly common in marshes and other freshwater habitats.
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.
Arcellinid testate amoebae or Arcellinida, Arcellacean or lobose testate amoebae are single-celled protists partially enclosed in a simple test (shell).
A protist is any eukaryotic organism that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor, the exclusion of other eukaryotes means that protists do not form a natural group, or clade. Therefore, some protists may be more closely related to animals, plants, or fungi than they are to other protists; however, like algae, invertebrates, or protozoans, the grouping is used for convenience. The study of protists is termed protistology.
Protozoa is an informal term for a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Historically, protozoans were regarded as "one-celled animals", because they often possess animal-like behaviours, such as motility and predation, and lack a cell wall, as found in plants and many algae. Although the traditional practice of grouping protozoa with animals is no longer considered valid, the term continues to be used in a loose way to describe single-celled protists that feed by heterotrophy. Some examples of protozoa are Amoeba, Paramecium, Euglena and Trypanosoma.
Arcella is a genus of testate amoebae in the order Arcellinida, usually found in freshwaters and mosses, and rarely in soils. A key characteristic of Arcella is the circular test with a hole on its center from where finger-like pseudopods emerge. It is one of the largest testacean genera.
Discorbacea, Discorboidea in recent taxonomies, is a superfamily of foraminifera,, with a range extending from the Middle Triassic to the present, characterized by chambers arranged in a low trochspiral; an umbilical or interiomarginal aperture, with or without supplementary apertures; and a wall structure that is optically radial.
Clavulina is a genus of aggulinated benthic foraminiferans with an elongate test. The early stage is triserial and triangular in section, the later stage uniserial and rectilinear, with angular to rounded section. In some species agglutinated walls have considerable calcareous cement. Septa are secondarily doubled as a result of imperforate floors, which are added as new chambers are formed. Walls contain fine bifurcating canaliculi within, openings of which are sealed internally by an inner organic lining, and externally by the imperforate surface layer of the wall. The aperture is interiomarginal in the early triserial stage, terminal and rounded in the adult.
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".
An amoeba, often called an amoeboid, is a type of cell or unicellular organism which has 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.
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