Testate amoebae

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Shell of Difflugia acuminata: an agglutinated test made up of mineral particles glued together with secretions from within the cell Difflugia acuminata.jpg
Shell of Difflugia acuminata : an agglutinated test made up of mineral particles glued together with secretions from within the cell

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.

Contents

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]

Taxonomy and classification

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]

Order Arcellinida

Family Arcellidae
  • Arcella - Ehrenberg 1832
  • Antarcella - Deflandre 1928
Family Netzeliidae
  • Netzelia - Ogden 1979
Family Hyalospheniidae
  • Quadrulella - Cockerell 1909
  • Hyalosphenia - Stein 1859
  • Alocodera - Jung 1942
  • Apodera - Loeblich & Tappan 1961
  • Certesella - Loeblich & Tappan 1961
  • Porosia - Jung 1942
  • Nebela - Leidy 1874
  • Padaungiella - Lara & Todorov 2012
Family Microchlamyiidae
  • Microchlamys - Cockerell 1911
  • Spumochlamys - Kudryavtsev & Hausmann 2007
Family Plagiopyxidae
  • Bullinularia - Deflandre 1953
  • Geoplagiopyxis - Chardez 1961
  • Protoplagiopyxis - Bonnet 1962
  • Paracentropyxis - Bonnet 1960
  • Plagiopyxis - Penard 1910
  • Hoogenraadia - Gauthier-Lievre & Thomas 1958
  • Planhoogenraadia - Bonnet 1977
Family Cryptodifflugiidae
Family Microcoryciidae
  • Amphizonella - Greeff 1866
  • Diplochlamys - Greeff 1888
  • Microcorycia - Cockerell 1911
  • Penardochlamys - Deflandre 1953
  • Zonomyxa - Nusslin 1882
  • Parmulina - Penard 1902
Family Phryganellidae
  • Phryganella - Penard 1902
Family Lamtopyxidae
  • Lamtopyxis - Bonnet 1974
Family Distomatopyxidae
  • Distomatopyxis - Bonnet 1964
Family Paraquadrulidae
  • Paraquadrula - Deflandre 1932
  • Lamtoquadrula - Bonnet 1974
Family Centropyxidae
  • Centropyxis - Stein 1857
  • Proplagiopyxis - Schonborn 1964
Family Trigonopyxidae
  • Trigonopyxis - Penard 1912
  • Cyclopyxis - Deflandre 1929
  • Geopyxella - Bonnet & Thomas 1955
  • Cornuapyxis - Couteaux and Chardez 1981
Incertae sedis
  • Argynnia - Vucetich 1974
  • Awerintzewia - Schouteden 1906
  • Cucurbitella - Penard 1902
  • Difflugia - Leclerc 1815
  • Geamphorella - Bonnet 1959
  • Heleopera - Leidy 1879
  • Jungia - Loeblich and Tappan 1961
  • Lagenodifflugia - Medioli & Scott 1983
  • Leptochlamys - West 1901
  • Lesquereusia - Schlumberger 1845
  • Maghrebia - Gauthier-Lievre & Thomas 1960
  • Mediolus - Patterson 2014
  • Microquadrula - Golemansky 1968
  • Oopyxis - Jung 1942
  • Pentagonia - Gauthier-Lievre & Thomas 1960
  • Physochila - Jung 1942
  • Pomoriella - Golemansky 1970
  • Pontigulasia - Rhumbler 1896
  • Protocucurbitella - Gauthier-Lievre & Thomas 1960
  • Pseudawerintzewia - Bonnet 1959
  • Pseudonebela - Gauthier-Lievre 1953
  • Pyxidicula - Ehrenberg 1838
  • Schoenbornia - Decloitre 1964
  • Schwabia - Jung 1942
  • Sexangularia - Awerintzew 1906
  • Suiadifflugia - Green 1975
  • Zivkovicia - Ogden 1987
  • Ellipsopyxis - Bonnet 1965
  • Ellipsopyxella - Bonnet 1975

Order Euglyphida

Other Cercozoa

Order Stramenopila

Unclassified testate amoebae

  • Paramphitrema - Valkanov 1970

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 ArcellinaAmphizonella - Arcella - Microchlamys - Microcorycia - Spumochlamys
DiffluginaBullinularia - Centropyxis - Difflugia - Distomatopyxis - Heleopera - Hyalosphenia - Lesquereusia - Nebela - Paraquadrula - Pontigulasia -

Plagiopyxis - Quadrulella - Trigonopyxis

Phryganellina Cryptodifflugia - Phryganella - Wailesella
DiscoseaHimatismenida Cochliopodium
SAR Supergroup StramenopilaLabyrinthulomycetesAmphitremida Amphitrema - Archerella
RhizariaCercozoaThecofiloseaCryomonadidaRhizaspididaeCapsellina - Rhizaspis - Rhogostoma
VentricleftidaVentrifissura - Verrucomonas
ImbricateaSilicofilosea Euglyphida Euglyphidae Euglypha - Scutiglypha
AssulinidaeAssulina - Placocista - Valkanovia
TrinematidaeCorythion - Playfairina - Puytoracia - Trinema
CyphoderidaeCampascus - Corythionella - Cyphoderia - Messemvriella - Pseudocorythion - Schaudinnula.
PaulinellidaeOvulinata - 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]

Notes

The Thecamoebida (Amoebozoa), with the genus Thecamoeba , despite their name, do not have tests.

Euglyphid testate amoebae are closely related to the Foraminifera. [9]

Related Research Articles

<span class="mw-page-title-main">Lobosa</span> Phylum of protozoans

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.

<span class="mw-page-title-main">Euglyphida</span> Order of single-celled organisms

The euglyphids are a prominent group of filose amoebae that produce shells or tests that in most described species is reinforced by siliceous scales, plates, and sometimes spines, but this reinforcement is absent in other species. 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.

<span class="mw-page-title-main">Amoebidae</span> Family of protozoans

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.

<i>Difflugia</i> Large genus of protists

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 biogenic elements and are thus commonly referred to as testate amoebae or shelled amoebae. Difflugia are particularly common in marshes and other freshwater habitats.

<span class="mw-page-title-main">Tubulinea</span> Class of protozoans

The Tubulinea are a major grouping of Amoebozoa, including most of the more familiar amoebae genera like Amoeba, Arcella, Difflugia and Hartmannella.

<span class="mw-page-title-main">Discosea</span> Class of amoebae

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.

<span class="mw-page-title-main">Arcellinida</span> Order of Amoebozoa

Arcellinid testate amoebae or Arcellinida, Arcellacean or lobose testate amoebae are single-celled protists partially enclosed in a simple test (shell).

<i>Arcella</i> Genus of protozoa

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.

<i>Apodera</i> Genus of shelled amoebae

Apodera is a genus of amoeboid protists belonging to the family Hyalospheniidae, a group of shelled amoebae. Their shells, or tests, are lageniform with a clear constriction that separates the neck from the body.

<i>Cryptodifflugia</i> Genus of testate amoebae

Cryptodifflugia is a genus of arcellinid testate amoebae. It contains all the species previously grouped as the genus Difflugiella, which is now a synonym of Cryptodifflugia.

<span class="mw-page-title-main">Amoeba</span> Cellular body type

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.

<span class="mw-page-title-main">Hyalospheniidae</span> Family of testate amoebae

Hyalospheniidae is a family of arcellinid testate amoebae and the sole family of the infraorder Hyalospheniformes. Commonly referred to as "hyalospheniids", these lobose amoebae are characterized by their ability to generate a shell composed of either organic matter or siliceous particles that may be recycled from euglyphid amoebae. They inhabit soil or freshwater habitats, and are abundant on Sphagnum mosses.

<i>Nebela</i> Genus of testate amoebae

Nebela is a diverse genus of testate amoebae of cosmopolitan distribution, belonging to the family Hyalospheniidae. They are "prey agglutinated" or "kleptosquamic" organisms, meaning they take the inorganic plates from their prey to construct their test.

<i>Certesella</i> Genus of testate amoebae

Certesella is a genus of testate amoebae belonging to the family Hyalospheniidae. It is characterized by a test that presents two symmetrical holes near the opening, and by the presence of internal teeth within the test. It contains four species previously assigned to Nebela, as well as one species discovered in 2021.

<span class="mw-page-title-main">Cryptodifflugiidae</span> Family of testate amoebae

Cryptodifflugiidae is a family of arcellinid testate amoebae.

<i>Meisterfeldia</i> Genus of testate amoebae

Meisterfeldia is a genus of arcellinid testate amoebae erected in 2016 that unites several new species as well as old species previously found in the genus Cryptodifflugia. Five of the newly described species were first found and isolated from subarctic tundra soil surrounding a river near Chokurdakh, Russia, while the last one, described in 2021, was recovered from tree hollows in Moscow.

<i>Planocarina</i> Genus of testate amoebae

Planocarina is a genus of arcellinid testate amoebae belonging to the family Hyalospheniidae. It was created in 2016 to agglutinate a clade of species that were previously assigned to the paraphyletic genus Nebela. All species of Planocarina have a compressed keel surrounding the posterior part of their shell. It is the sister group of Alabasta.

<i>Alabasta</i> Genus of testate amoebae

Alabasta is a genus of arcellinid testate amoebae belonging to the family Hyalospheniidae. It contains species with an elongated test and a strongly curved "pseudostome" with a flare and a notch in narrow view. These species previously belonged to the genus Nebela, but were later found to be a distinct monophyletic group different from Nebela. It is the sister group to Planocarina.

<span class="mw-page-title-main">Corycidia</span> Group of amoebae

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.

<i>Cryptodifflugia operculata</i> Species of ameobozoans

Cryptodifflugia operculata is an amoebozoan in the Tubulinea class. They are testate amoebae due to the presence of a test that encapsulates its pseudopods. C. operculata's taxonomy has been categorized throughout the years, as well as its morphology. They are found mainly in freshwater, however they also can be found in soil. C. operculata is a predator that can consume prey larger than themselves, and is capable of forming packs and hunting.

References

  1. B., Scott, D. (2001). Monitoring in coastal environments using Foraminifera and Thecamoebian indicators. Medioli, F. S., Schafer, Charles T. Cambridge: Cambridge University Press. ISBN   978-0521561730. OCLC   70724931.{{cite book}}: CS1 maint: multiple names: authors list (link)
  2. 1 2 Kosakyan, Anush; Gomaa, Fatma; Lara, Enrique; Lahr, Daniel J.G. (2016). "Current and future perspectives on the systematics, taxonomy and nomenclature of testate amoebae". European Journal of Protistology. 55 (Pt B): 105–117. doi:10.1016/j.ejop.2016.02.001. hdl: 11380/1302106 . PMID   27004416.
  3. Sigala, Itzel; Lozano-García, Socorro; Escobar, Jaime; Pérez, Liseth; Gallegos-Neyra, Elvia (2016-06-28). "Testate Amoebae (Amebozoa: Arcellinida) in Tropical Lakes of Central Mexico". Revista de Biología Tropical. 64 (1): 377–397. doi: 10.15517/rbt.v64i1.18004 . ISSN   2215-2075. PMID   28862826.
  4. Mitchell, Edward A. D.; Charman, Daniel J.; Warner, Barry G. (2008-08-01). "Testate amoebae analysis in ecological and paleoecological studies of wetlands: past, present and future" (PDF). Biodiversity and Conservation. 17 (9): 2115–2137. doi:10.1007/s10531-007-9221-3. ISSN   0960-3115. S2CID   27912982.
  5. Ralf Meisterfeld: Arcellinida, In: John J. Lee, Gordon F. Leedale, Phyllis Bradbury (Hrsg.): Illustrated Guide to the Protozoa, 2nd Edition. Vol. 2, Society of Protozoologists, Lawrence, Kansas 2000, ISBN   1-891276-23-9, pp. 827-860
  6. Ralf Meisterfeld: Testate amoebae with filopodia , In: John J. Lee, Gordon F. Leedale, Phyllis Bradbury (Hrsg.): The Illustrated Guide to the Protozoa, 2nd Edition. Vol. 2, Society of Protozoologists, Lawrence, Kansas 2000, ISBN   1-891276-23-9, pp. 1054-1084
  7. 1 2 3 Adl, Sina M.; Simpson, Alastair G. B.; Lane, Christopher E.; Lukeš, Julius; Bass, David; Bowser, Samuel S.; Brown, Matthew W.; Burki, Fabien; Dunthorn, Micah (2012-09-01). "The Revised Classification of Eukaryotes". Journal of Eukaryotic Microbiology. 59 (5): 429–514. doi:10.1111/j.1550-7408.2012.00644.x. ISSN   1550-7408. PMC   3483872 . PMID   23020233.
  8. Ruggiero, Michael A.; Gordon, Dennis P.; Orrell, Thomas M.; Bailly, Nicolas; Bourgoin, Thierry; Brusca, Richard C.; Cavalier-Smith, Thomas; Guiry, Michael D.; Kirk, Paul M. (2015-04-29). "A Higher Level Classification of All Living Organisms". PLOS ONE. 10 (4): e0119248. Bibcode:2015PLoSO..1019248R. doi: 10.1371/journal.pone.0119248 . ISSN   1932-6203. PMC   4418965 . PMID   25923521.
  9. Testate amoebae as environmental indicators (PDF)

Bibliography