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Arcellinida Temporal range: Tonian–Present Pha. Proterozoic Archean Had.
Arcella sp. test
Scientific classification
Domain:	Eukaryota
Phylum:	Amoebozoa
Class:	Tubulinea
Superorder:	Eulobosia
Order:	Arcellinida Kent 1880

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

Arcellinid testate amoebae are commonly found in soils, leaf litter, peat bogs and near/in fresh water. They use their pseudopodia, a temporary cell extension, for moving and taking in food. Like most amoebae, they are generally believed to reproduce asexually via binary fission. However a recent review suggests that sexual recombination may be the rule rather than the exception in amoeboid protists in general, including the Arcellinid testate amoebae.

Morphology

Arcellinida always have a shell or testa. The tests lie outside the cell membrane and consist of organic or mineral materials that are either secreted or incorporate external particles. The testa has a single main opening.

Simple tests are made by secretion (autogenous tests), agglutination of foreign material (xenogenous tests), or sometimes a combination of both. Past environmental changes can be determined by analysing the composition of fossil tests, including the reconstruction of past climate change. Testate amoebae species have been used to reconstruct hydrological changes over the late Holocene, as a result of individual species possessing a narrow tolerance for ecohydrological conditions such as water-table depth or pH.

Evolutionary history

Fossils of arcellinid testate amoebae date back to the Tonian stage of the Proterozoic, around 789-759 million years ago. The fossils indicate that by 730 million years ago, arcellinids had already diversified into major lineages.

Testate amoebae are theorized to be mostly polyphyletic (coming from more than one ancestral type), but testaceafilosea, one group of testate amoebae, are theorized to be monophyletic. Ancient tests of terrestrial fauna are commonly found in fossilized amber, although mid-Cretaceous testate amoeba (i.e., Diffligia, Cucurbitella) have been found in ancient lake sediments. It is likely that the group has evolved minimally over the course of the Phanerozoic.

Classification

The classification of Arcellinida, as of 2019:

• Suborder Phryganellina Bovee 1985
  • Family Phryganellidae Jung 1942
    • Phryganella Penard 1902
  • Family Cryptodifflugiidae Jung 1942
    • ?Prantlitina Vasicek & Ruzicka 1957
    • ?Pseudocucurbitella Gauthier-Lievre & Thomas 1960
    • ?Pseudowailesella Sudzuki 1979
    • Cryptodifflugia Penard 1890
    • Meisterfeldia Bobrov 2016
    • Wailesella Deflandre 1928
• Suborder Organoconcha Lahr et al. 2019
  • Family Microchlamyiidae Ogden 1985
    • Microchlamys Cockerell 1911
    • Pyxidicula Ehrenberg 1838 non Ehrenberg 1834 non Strelnikova & Nikolajev 1986
    • Spumochlamys Kudryavtsev & Hausmann 2007
• Suborder Glutinoconcha Lahr et al. 2019
  • Infraorder Volnustoma Lahr et al. 2019
    • Family Heleoperidae Jung 1942
      • Awerintzewia Schouteden 1906
      • Heleopera Leidy 1879
  • Infraorder Hyalospheniformes Lahr et al. 2019
    • Family Hyalospheniidae Schulze, 1977
      • ?Apolimia Korganova 1987
      • ?Deflandria Jung 1942
      • ?Heleoporella Couteaux 1978
      • ?Leidyella Jung 1942
      • ?Metaheleopera Bartos 1954
      • ?Marsipos Medioli et al. 1990
      • ?Paranebela Jung 1942
      • ?Penardiella Kahl 1930
      • ?Pseudogeamphorella Décloitre 1964 nomen nudum
      • ?Pseudohyalosphenia Stepanek 1967 nomen nudum
      • ?Pterygia Jung 1942 non Roeding 1798 non Link 1807 non Laporte 1832
      • ?Schaudinnia Jung 1942 non Schulze 1900
      • ?Umbonaria Jung 1942
      • Alabasta Duckert et al. 2018
      • Alocodera Jung 1942a
      • Apodera Loeblich & Tappan 1961
      • Certesella Loeblich & Tappan 1961
      • Cornutheca Kosakyan et al. 2016
      • Gibbocarina Kosakyan et al. 2016
      • Hyalosphenia Stein 1859
      • Longinebela Kosakyan et al. 2016
      • Mrabella Kosakyan et al. 2016
      • Nebela Leidy 1875
      • Padaungiella Lara & Todorov 2012
      • Planocarina Kosakyan et al. 2016
      • Porosia Jung 1942
      • Quadrulella Cockerell 1909
  • Infraorder Excentrostoma Lahr et al. 2019
    • ?Centropyxiella Valkanov 1970
    • ?Oopyxis Jung 1942
    • Family Centropyxidae
      • Armipyxis Dekhtiar 2009
      • Centropyxis Stein 1857
      • Conicocassis Nasser & Patterson 2015
      • Proplagiopyxis Schönborn 1964
    • Family Plagiopyxidae
      • Bullinularia Deflandre 1953
      • Geoplagiopyxis Chardez 1961
      • Hoogenraadia GauthierLievre & Thomas 1958
      • Paracentropyxis Bonnet 1960
      • Plagiopyxis Penard 1910
      • Planhoogenraadia Bonnet 1977
      • Protoplagiopyxis Bonnet 1962
  • Infraorder Longithecina Lahr et al. 2019
    • Family Lesquereusiidae
      • Fabalesquereusia Snegovaya & Alekperov 2005
      • Lesquereusia Schlumberger 1845
      • Microquadrula Golemansky 1968
      • Paraquadrula Deflandre 1932
      • Pomoriella Golemansky 1970
    • Family Difflugiidae Wallich 1864
      • ?Lagenodifflugia Medioli & Scott 1983
      • ?Maghrebia Gauthier-Lievre & Thomas 1958
      • ?Pentagonina Bovee & Jahn 1974
      • ?Pseudopontigulasia Oye 1956
      • ?Sexangularia Awerintzew 1906
      • ?Suiadifflugia Green 1975
      • ?Zivkovicia Ogden 1987
      • Armatodifflugia Snegovaya & Alekperov 2005
      • Difflugia Leclere 1815 ex Lamarck 1816
      • Mediolus Patterson 2014
      • Nabranella Snegovaya & Alekperov 2009
      • Pelecyamoeba Snegovaya & Alekperov 2005
      • Pontigulasia Rhumbler 1895
      • Pseudonebela Gauthier-Lievre 1954 non Schönborn 1964
  • Infraorder Sphaerothecina Kosakyan et al. 2016
    • ?Cornuapyxis Couteaux & Chardez 1981
    • ?Cucurbitella Penard 1902 non Walpers 1846
    • ?Ellipsopyxella Bonnet 1975
    • ?Ellipsopyxis Bonnet 1965
    • ?Protocucurbitella Gauthier-Lievre & Thomas 1960 non Naumov
    • Family Distomatopyxidae Bonnet 1970
      • Distomatopyxis Bonnet 1970
    • Family Lamtopyxidae Bonnet 1974
      • Lamtopyxis Bonnet 1974
    • Family Netzeliidae Kosakyan et al. 2016
      • Cyclopyxis Bonnet 1953
      • Netzelia Ogden 1979
    • Family Arcellidae Ehrenberg 1832
      • Antarcella Ehrenberg 1838
      • Arcella Deflandre 1928
    • Trigonopyxidae Loeblich 1964
      • Geopyxella Bonnet & Thomas 1955
      • Trigonopyxis Penard 1912

Arcellinida incertae sedis:

• Argynnia Vucetich, 1974
• Awerintzewia Schouteden, 1906
• Geamphorella Bonnet, 1959
• Jungia Loeblich & Tappan 1961
• Lagenodifflugia Medioli & Scott, 1983
• Lamtoquadrula Bonnet 1975
• Leptochlamys West 1901
• Maghrebia Gauthier-Lievre & Thomas, 1960
• Ochros Medioli et al. 1990
• †Palaeoleptochlamys Strullu-Derrien et al. 2019
• Physochila Jung 1942
• Pseudawerintzewia Bonnet 1959
• Sacculus Medioli et al. 1990 non Gosse 1851 non Hirase 1927 non Neviani 1930
• Schoenbornia Decloitre 1964
• Swabia
• Family Bipseudostomatidae Snegovaya & Alekperov 2005
  • Bipseudostomatella Snegovaya & Alekperov 2005
  • Gomocollariella Snegovaya & Alekperov 2005
• Family Mississippiellidae Huddleston & Haman 1985
  • Mississippiella Haman 1982
• Family Shamkiriidae Snegovaya & Alekperov 2005
  • Shamkiriella Snegovaya & Alekperov 2005

References

1. ^ http://www.microscopy-uk.org.uk/mag/artjun03/gsamoebae.html Testate amoebae, peat bogs and past climates. accessed 16 march 2007
2. Identification key for holocene lacustrine arcellacean (thecamoebian) taxa Archived December 4, 2008, at the Wayback Machine
3. Lara E, Heger TJ, Ekelund F, Lamentowicz M, Mitchell EA (April 2008). "Ribosomal RNA genes challenge the monophyly of the Hyalospheniidae (Amoebozoa: Arcellinida)". Protist. 159 (2): 165–76. doi:10.1016/j.protis.2007.09.003. PMID 18023614.
4. Lahr DJ, Parfrey LW, Mitchell EA, Katz LA, Lara E (2011). "The chastity of amoebae: re-evaluating evidence for sex in amoeboid organisms". Proceedings of the Royal Society B: Biological Sciences. 278 (1715): 2081–2090. doi:10.1098/rspb.2011.0289. PMC 3107637. PMID 21429931.
5. ^ Adl SM, Simpson AG, Lane CE, Lukeš J, Bass D, Bowser SS, Brown MW, Burki F, Dunthorn M, Hampl V, Heiss A, Hoppenrath M, Lara E, le Gall L, Lynn DH, McManus H, Mitchell EA, Mozley-Stanridge SE, Parfrey LW, Pawlowski J, Rueckert S, Shadwick L, Schoch CL, Smirnov A, Spiegel FW (2012). "The Revised Classification of Eukaryotes". Journal of Eukaryotic Microbiology. 59 (5): 429–514. doi:10.1111/j.1550-7408.2012.00644.x. PMC 3483872. PMID 23020233.
6. Woodland, Wendy, A. (1998). "Quantitative estimates of water tables and soil moisture in Holocene peatlands from testate amoebae". The Holocene. 8 (3): 261–273. doi:10.1191/095968398667004497. S2CID 128897334.{{cite journal}}: CS1 maint: multiple names: authors list (link)
7. Lahr, Daniel J.G.; Kosakyan, Anush; Lara, Enrique; Mitchell, Edward A.D.; Morais, Luana; Porfirio-Sousa, Alfredo L.; Ribeiro, Giulia M.; Tice, Alexander K.; Pánek, Tomáš; Kang, Seungho; Brown, Matthew W. (2019-03-18). "Phylogenomics and Morphological Reconstruction of Arcellinida Testate Amoebae Highlight Diversity of Microbial Eukaryotes in the Neoproterozoic". Current Biology. 29 (6): 991–1001.e3. doi:10.1016/j.cub.2019.01.078. hdl:10261/240246. ISSN 0960-9822. PMID 30827918. S2CID 72333352.
8. Schmidt, A.R.; Ragazzi, E.; Coppellotti, O.; Roghi, G. (2006). "A microworld in Triassic amber". Nature. 444 (7121): 835. doi:10.1038/444835a. PMID 17167469.
9. Hengstum, Van; Reinhardt, E.G.; Medioli, F.S.; Grocke, D.R. (2007). "Exceptionally preserved late albian (Cretaceous) Arcellaceans (Thecamoebians) from the Dakota Formation near Lincoln, Nebraska" (PDF). Journal of Foraminiferal Research. 37 (4): 300–308. doi:10.2113/gsjfr.37.4.300.
10. ^ Lahr, Daniel J.G.; Kosakyan, Anush; Lara, Enrique; Mitchell, Edward A.D.; Morais, Luana; Porfirio-Sousa, Alfredo L.; Ribeiro, Giulia M.; Pánek, Tomáš; Kang, Seungho; Brown, Matthew W. (2019). "Phylogenomics and Morphological Reconstruction of Arcellinida Testate Amoebae Highlight Diversity of Microbial Eukaryotes in the Neoproterozoic". Current Biology. 29 (6): 991–1001. doi:10.1016/j.cub.2019.01.078. hdl:11380/1301486. PMID 30827918.
11. González-Miguéns, Rubén; Todorov, Milcho; Blandenier, Quentin; Duckert, Clément; Porfirio-Sousa, Alfredo L.; Ribeiro, Giulia M.; Ramos, Diana; Lahr, Daniel J.G.; Buckley, David; Lara, Enrique (2022). "Deconstructing Difflugia: The tangled evolution of lobose testate amoebae shells (Amoebozoa: Arcellinida) illustrates the importance of convergent evolution in protist phylogeny". Molecular Phylogenetics and Evolution. 175: 107557. doi:10.1016/j.ympev.2022.107557. hdl:10261/281619. ISSN 1055-7903. PMID 35777650.
12. 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

External links

• ISTAR, the International Society for Testate Amoeba Research
• Tree of Life: Arcellinida
• An article by G. T. Swindles
• Protozoa: The testate amoebae
• BioImages: The Virtual Field-Guide (UK) - TESTACEA (Testate amoebae)
• Pictures of testate amoebae
• The Cushman Foundation Thecamoebian Page

• Amoebozoa orders
• Tubulinea