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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.

Testate amoebae (thecamoebians, formally Testacea or Thecamoeba) are a polyphyletic group of unicellular ameboid protists, which differ from naked amoebae in the presence of a test that partially encloses the cell, from which the pseudopodia emerge, and that provides the cell with shelter from predators and environmental conditions.

The test of some species is made up of particles of sediment where the amoebae live, which are agglutinated together by secretions from within the cell, whereas in other cases the test is produced entirely by the amoeba and may be organic, siliceous or calcareous depending on the species.

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.[1]

Testate amoebae can be found in most freshwater environments, including lakes, rivers, cenotes[2], 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. This, as well as 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.[3]

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)[4], and the filose Euglyphida (within the SAR supergroup)[5], although there are smaller groups that also include other testate amoebae.[6]

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

    Cryptodifflugia Penard 1890
    Wailesella Deflandre 1928

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

Family Euglyphidae

    Euglypha Dujardin, 1841
    Scutiglypha Foissner and Schiller, 2001

Family Trinematidae

    Trinema Dujardin 1841
    Corythion Taranek 1881
    Puytoracia Bonnet, 1970
    Playfairina Thomas 1961
    Pileolus Coûteaux and Chardez 1962

Family Sphenoderiidae

    Sphenoderia Schlumberger 1845
    Trachelocorythion Bonnet 1979
    Deharvengia Bonnet 1979

Family Assulinidae

    Assulina Leidy 1879
    Placocista Leidy 1879
    Valkanovia Tappan 1966

Family Cyphoderiidae

    Cyphoderia Schlumberger 1845
    Corythionella Golemansky 1970
    Campascus Leidy 1879
    Messemvriella Golemansky 1973
    Pseudocorythion Valkanov 1970
    Schaudinnula Awerintzev 1907

Family Paulinellidae

    Paulinella Lauterborn 1895
    Ovulinata Anderson, Rogerson & Hannah 1996
    Micropyxidiella Tarnawski & Lara 2015

Incertae sedis

    Pareuglypha Penard 1902
    Ampullataria Van Oye 1956
    Euglyphidion Bonnet 1960
    Heteroglypha Thomas and Gauthier-Lièvre 1959
    Tracheleuglypha Deflandre 1928

Other Cercozoa

Family Chlamydophryidae

    Capsellina Penard 1909
    Lecythium Hertwig and Lesser 1876
    Rhogostoma Belar 1921
    Rhizaspis Skuja 1948
    Chlamydophrys Cienkowsky 1876
    Clypeolina Cienkowsky 1877
    Penardeugenia Deflandre 1953
    Leptochlamydophrys Belar 1921
    Diaphoropodon Archer 1869

Family Pseudodifflugiidae

    Pseudodifflugia Schlumberger 1845

Family Psammonobiotidae

    Alepiella Golemansky 1970
    Centropyxiella Valkanov 1970
    Chardezia Golemansky 1970
    Edaphonobiotus Schönborn, Foissner & Meisterfeld 1983
    Micramphora Valkanov 1970
    Micropsammella Golemansky 1970
    Nadinella Penard 1902
    Ogdeniella Golemanski 1982
    Psammonobiotus Golemansky 1970
    Propsammonobiotus Golemansky 1991

Incertae sedis

    Feuerbornia Jung, 1942
    Frenzelina Penard 1902
    Lesquerella Chardez and Thomas, 1980
    Matsakision Bonnet, 1967
    Rhumbleriella Golemansky 1970

Order Stramenopila

Family Amphitremidae

    Amphitrema Archer 1869
    Archerella Loeblich and Tappan, 1961

Family Diplophryidae

    Diplophrys Barker 1868

Family Amphifilidae

    Amphifila Anderson and Cavalier-Smith 2012

Family Sorodiplophryidae

    Sorodiplophris Dykstra and Olive 1975

Unclassified testate amoebae

    Paramphitrema Valkanov 1970


The following table includes a few examples of testate amoebae genera, and reflects the classification by Adl et al. (2012)[6], 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[7], 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.[6]

Amoebozoa Tubulinea Arcellinida Arcellina Amphizonella - Arcella - Microchlamys - Microcorycia - Spumochlamys  
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).[1]

Notes

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

Euglyphid testate amoebae are closely related to the Foraminifera.[8]

See also

  • Microworld - World of ameboid organisms - A database of both testate and naked amoebae with over 6,700 microphotographs and videos and over 1,700 species descriptions, as well as dichotomous and visual keys for identification.

References

  1. ^ a b Kosakyan, Anush; Gomaa, Fatma; Lara, Enrique; Lahr, Daniel J.G. "Current and future perspectives on the systematics, taxonomy and nomenclature of testate amoebae". European Journal of Protistology. 55: 105–117. doi:10.1016/j.ejop.2016.02.001.
  2. ^ 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.
  3. ^ 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". Biodiversity and Conservation. 17 (9): 2115–2137. doi:10.1007/s10531-007-9221-3. ISSN 0960-3115.
  4. ^ 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
  5. ^ 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
  6. ^ a b c 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.
  7. ^ 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. doi:10.1371/journal.pone.0119248. ISSN 1932-6203.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  8. ^ Testate amoebae as environmental indicators (PDF)

Bibliography

  • Medioli, F.S.; Scott, D.B.; Collins, E.; Asioli, S.; Reinhardt, E.G. (1999). The thecamoebian bibliography. Palaeontologia Electronica, 3: 1-161, [1].
  • Medioli, F.S.; Bonnet, L.; Scott, D.B.; Medioli, B.E. (2003). The thecamoebian bibliography: 2nd edition. Palaeontologia Electronica, 61: 1-107, [2].