Tubulinea

Tubulinea
	Amoeba proteus
Scientific classification
Domain:	Eukaryota
Phylum:	Amoebozoa
Subphylum:	Lobosa
Class:	Tubulinea; Smirnov et al. 2005
Subclasses & Orders
	Corycidia Trichosida; ; Echinamoebia Echinamoebida; ; Elardia Leptomyxida; Euamoebida; Arcellinida; ;

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

Characteristics

During locomotion most Tubulinea have a roughly cylindrical form or produce numerous cylindrical pseudopods. Each cylinder advances by a single central stream of cytoplasm, granular in appearance, and has no subpseudopodia. This distinguishes them from other amoeboid groups, although in some members this is not the normal type of locomotion.

**Representation of a tubulinid**
Ectoplasm
Endoplasm
Lobopodia, a pseudopod or arm-like projection made of cytoplasm
Phagocytic vacuole
Mitochondrion, creates ATP (energy) for the cell (branched cristae)
Endosymbiotic bacterium
Triuret (nitrogen waste) crystal
Prey
Nucleus
Honeycomb lamina
Nucleolus
Endoplasmic reticulum, the transport network for molecules going to specific parts of the cell
Golgi apparatus, modifies proteins and sends them out of the cell
Lysosome, holds enzymes
Digestive vacuole
Lipid granule
Vesicle
Contractile vacuole, regulates the quantity of water inside a cell
Uroid

Classification

This class was anticipated by some biologists such as Jahn, who grouped all amoebae with granular pseudopodia together,^[1] but most split the lobose amoebae into testate Testacealobosia and naked Gymnamoebia. These latter are polyphyletic, but molecular trees by Bolivar et al.^[2] identified a core monophyletic subgroup. Subsequent studies showed the testate lobose amoebae belong to the same group, which was thus renamed Lobosea sensu stricto^[3] or Tubulinea.^[4]

Taxonomy

The class Tubulinea, as of 2022, is classified into three major groups: Corycida, Echinamoebida and Elardia. The most taxonomically abundant group is Elardia, which contains the testate amoebae of Arcellinida and the naked amoebae of orders Leptomyxida and Euamoebida.^[5]^[6]^[7]^[8]

Corycidia Kang et al. 2017
- Family Microcoryciidae de Saedeleer 1934
- Order Trichosida Moebius 1889
  - Family Trichosphaeriidae Sheehan & Banner 1973
Order Echinamoebida Cavalier-Smith 2004 em. 2011
- Family Vermamoebidae Cavalier-Smith & Smirnov 2011
- Family Echinamoebidae Page 1975
Elardia Kang et al. 2017
- Order Leptomyxida Pussard & Pons 1976 em. Page 1987
  - Family Flabellulidae Bovee & Jahn, 1967 ex Bovee 1970
  - Family Gephyramoebidae Pussard & Pons 1976
  - Family Leptomyxidae Goodey 1915
  - Family Rhizamoebidae Smirnov et al. 2016
- Superorder Eulobosia Cavalier-Smith 2016
  - Order Euamoebida Lepşi 1960 em. Cavalier-Smith 2016
    - Family Nolandellidae Cavalier-Smith 2011
    - Family Hartmannellidae Singh 1951 em. Smirnov et al. 2011
    - Family Amoebidae (Ehrenberg 1838) Page 1987
  - Order Arcellinida Kent 1880
    - Suborder Phryganellina Bovee 1985
      - Family Phryganellidae Jung 1942
      - Family Cryptodifflugiidae Jung 1942
    - Suborder Organoconcha Kosakyan et al. 2016
      - Family Microchlamyiidae Ogden 1985
    - Suborder Glutinoconcha Kosakyan et al. 2016
      - Infraorder Volnustoma Kosakyan et al. 2016
        Family Heleoperidae Jung 1942
      - Infraorder Hyalospheniformes Lahr et al. 2019
        Family Hyalospheniidae Schultze 1877 emend. Kosakyan & Lara 2012 [Nebelidae Taranek 1882]
      - Infraorder Excentrostoma Lahr et al. 2019
        Family Centropyxidae Jung 1942 [incl. Plagiopyxidae]
      - Infraorder Cylindrothecina González-Miguéns, Todorov, Blandenier, Porfirio-Sousa, Ribeiro, Ramos, Lahr & Lara 2022
        Family Cylindrifflugiidae González-Miguéns, Todorov, Blandenier, Porfirio-Sousa, Ribeiro, Ramos, Lahr & Lara 2022
      - Infraorder Longithecina Kosakyan et al. 2016
        Family Lesquereusiidae [Paraquadrulidae Deflandre 1953]
        
        Family Difflugiidae Wallich 1864
      - Infraorder Sphaerothecina Kosakyan et al. 2016
        Family Netzeliidae Kosakyan et al. 2016 [Cyclopyxidae Schönborn 1989]
        
        Family Arcellidae Ehrenberg 1832

Amoeba proteus
Chaos sp.
Arcella sp. test
Trichosphaerium sieboldi

References

^ Jahn; Bovee; Jahn (1979). How to know the protozoa. Wm. C. Brown Company Publishers. ISBN 0-697-04759-8.
^ Bolivar, I.; Fahrni, J.F.; Smirnov, A. & Pawlowski, J. (2001). "SSU rRNA-based Phylogenetic Position of the Genera Amoeba and Chaos (Lobosea, Gymnamoebia): The Origin of Gymnamoebae Revisited". Molecular Biology and Evolution. 18 (12): 2306–2314. doi:10.1093/oxfordjournals.molbev.a003777. PMID 11719580.
^ Cavalier-Smith, T.; Chao, E.-Y; Oates, B. (2004). "Molecular phylogeny of the Amoebozoa and evolutionary significance of the unikont Phalansterium". European Journal of Protistology. 40: 21–48. doi:10.1016/j.ejop.2003.10.001.
^ Smirnov, A.V.; Nassonova, E.S.; Berney, C.; Fahrni, J.; Bolivar, I. & Pawlowski, J. (2005). "Molecular phylogeny and classification of the lobose amoebae". Protist. 156 (2): 129–142. doi:10.1016/j.protis.2005.06.002. PMID 16171181.
^ Kang S, Tice AK, Spiegel FW, Silberman JD, Pánek T, Cepicka I, et al. (September 2017). "Between a Pod and a Hard Test: The Deep Evolution of Amoebae". Molecular Biology and Evolution. 34 (9): 2258–2270. doi:10.1093/molbev/msx162. PMC 5850466. PMID 28505375.
^ Adl, Sina M.; Bass, David; Lane, Christopher E.; Lukeš, Julius; Schoch, Conrad L.; Smirnov, Alexey; Agatha, Sabine; Berney, Cedric; Brown, Matthew W. (2018-09-26). "Revisions to the Classification, Nomenclature, and Diversity of Eukaryotes". Journal of Eukaryotic Microbiology. 66 (1): 4–119. doi:10.1111/jeu.12691. PMC 6492006. PMID 30257078.
^ 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.
^ 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.

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[1] Jahn; Bovee; Jahn (1979). How to know the protozoa. Wm. C. Brown Company Publishers. ISBN 0-697-04759-8.

[2] Bolivar, I.; Fahrni, J.F.; Smirnov, A. & Pawlowski, J. (2001). "SSU rRNA-based Phylogenetic Position of the Genera Amoeba and Chaos (Lobosea, Gymnamoebia): The Origin of Gymnamoebae Revisited". Molecular Biology and Evolution. 18 (12): 2306–2314. doi:10.1093/oxfordjournals.molbev.a003777. PMID 11719580.

[3] Cavalier-Smith, T.; Chao, E.-Y; Oates, B. (2004). "Molecular phylogeny of the Amoebozoa and evolutionary significance of the unikont Phalansterium". European Journal of Protistology. 40: 21–48. doi:10.1016/j.ejop.2003.10.001.

[4] Smirnov, A.V.; Nassonova, E.S.; Berney, C.; Fahrni, J.; Bolivar, I. & Pawlowski, J. (2005). "Molecular phylogeny and classification of the lobose amoebae". Protist. 156 (2): 129–142. doi:10.1016/j.protis.2005.06.002. PMID 16171181.

[5] Kang S, Tice AK, Spiegel FW, Silberman JD, Pánek T, Cepicka I, et al. (September 2017). "Between a Pod and a Hard Test: The Deep Evolution of Amoebae". Molecular Biology and Evolution. 34 (9): 2258–2270. doi:10.1093/molbev/msx162. PMC 5850466. PMID 28505375.

[Adl2018-6] Adl, Sina M.; Bass, David; Lane, Christopher E.; Lukeš, Julius; Schoch, Conrad L.; Smirnov, Alexey; Agatha, Sabine; Berney, Cedric; Brown, Matthew W. (2018-09-26). "Revisions to the Classification, Nomenclature, and Diversity of Eukaryotes". Journal of Eukaryotic Microbiology. 66 (1): 4–119. doi:10.1111/jeu.12691. PMC 6492006. PMID 30257078.

[Lahr_2019-7] 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.

[Arcellinida2022-8] 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.

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