Taxonomy of Cercozoa

Cercozoa (=Rhizopoda) is a phylum of protists.[1] It was first described in 1998 by Thomas Cavalier-Smith. The phylum Cercozoa belongs to a group called Rhizaria. No specific morphological or behavioral characteristics are observed in its members. This phylum exhibits great diversity. Its members are biciliate and/or amoeboid and generally possess filopodia. Most contain tubular cristae, and cysts are common. Kinetosomes are attached to the nucleus via the cytoskeleton. Microbodies and extrusomes are generally present.[2] This list shows the revisions made to Cavalier-Smith's Cercozoa classification and also shows Adl et al. classifications.

Taxonomy

Before the phylum was described, a similar group was named as phylum Rhizopoda instead. In 1997, Rhizopoda was assigned to a superphylum named Neosarcodina that also included another phylum, the Amoebozoa. Here is a scheme from Cavalier-Smith, 1997:[3]

In Cavalier-Smith, 1998, the phylum was proposed as a replacement for Rhizopoda. It was placed in infrakingdom Sarcomastigota along with the phyla Neomonada, Foraminifera and Amoebozoa. Order Spongomonadida was added to subphylum Monadofilosa and cristidiscoids were transferred to subphylum Choanozoa. The classification was this:[1]

In Cavalier-Smith, 2000, a new monotypic class Spongomonadea was named for the order Spongomonadida. Spongomonadea was placed in subphylum Reticulofilosa. Order Cryomonadida was added to class Sarcomonadea. Class Athalamea was used for the families Biomyxidae and Gymnophryidae in subphylum Reticulofilosa. Class Ramicristea was used in subphylum Monadofilosa and included the orders Heliomonadida and Commatiida:[4][5][6][7]

In Cavalier-Smith, 2002, a new subphylum named Endomyxa was introduced to contain the new class Ascetosporea and the class Phytomyxea in the phylum Cercozoa. Ascetosporea contained the orders Haplosporida and Paramyxida. Subclass Testaceafilosia was ranked as a class Testaceafilosea. Phylum Cercozoa was placed in a new infrakingdom named Rhizaria along with the phyla Apusozoa, Retaria and Heliozoa:[8]

In Cavalier-Smith and Chao, 2003, the subphyla Reticulofilosa and Monadofilosa were ranked as superclasses in a new subphylum Filosa. Order Heliomonadida was transferred to class Proteomyxidea, a new order Reticulosida was created to include the only family Gymnophryidae, now also containing family Biomyxidae members. A new order Metopiida was assigned to class Sarcomonadea. A new class named Thecofilosea was created to contain the order Cryomonadida and the new order Tectofilosida. A new class named Imbricatea was created to contain the orders Thaumatomonadida and Euglyphida. A new order Claustrosporida was assigned to class Ascetosporea. A new class named Gromiidea contained the only order Gromiida. Order Commatiida was not listed, and order Vampyrellida was assigned kingdom Protozoa incertae sedis.[7]

In Bass and Cavalier-Smith, 2004, a new order Metromonadida was placed as superclass Monadofilosa incertae sedis. Order Metopiida was removed from class Sarcomonadea and placed subphylum Filosa incertae sedis instead. Class Spongomonadea was transferred to superclass Monadofilosa:[9]

In Bass et al., 2005, genus Sainouron was classified as superclass Monadofilosa incertae sedis, genera Helkesimastix and Cercobodo were also classified as subphylum Filosa incertae sedis:[10]

In 2005, Adl et al. proposed a rankless hierarchy:[2]

In Cavalier-Smith 2007, the order Metromonadida was assigned to its own class Metromonadea directly under subphylum Filosa and class Phaeodarea and order Metopiida was added to superclass Monadofilosa:[11]

In Cavalier-Smith et al., 2008, the orders Pansomonadida and Metopiida and family Sainouridae was added to class Sarcomonadea and order Desmothoracida was added to class Proteomyxidea:[12]

In Bass et al., 2009, a new class named Granofilosea was added to subphylum Filosa and contained five orders: Leucodictyida, Limnofilida, Cryptofilida, Desmothoracida and Heliomonadida. A new order Paradinida was added to class Ascetosporea. Order Aconchulinida was added to class Proteomyxidea, a new family named Rhizoplasmidae was also added to the class as incertae sedis. Class Proteomyxidea was transferred to subphylum Endomyxa:[6]

In Howe et al., 2009, a new order Glissomonadida was also assigned to class Sarcomonadea and genus Pseudopirsonia was added as superclass Monadofilosa incertae sedis:[13]

In Cavalier-Smith et al., 2009 a new superfamily Sainouroidea was named to include the family Sainouridae and the new monotypic family Helkesimastigidae including the genus Helkesimastix. Superfamily Sainouroidea was placed in class Sarcomonadea:[14]

In Howe et al., 2011, order Spongomonadida, a new order Marimonadida that also included the genus Pseudopirsonia and a new incertae sedis family Nudifilidae was added to class Imbricatea. Genera Clautriavia and Discomonas were also added as Imbricatea incertae sedis. Phaeodarea was ranked as a subclass Phaeodaria and added to class Thecofilosea. Order Ebriida and a new order Ventricleftida was also added in class Thecofilosea. Order Metopiida was transferred to class Metromonadea. Class Metromonadea was assigned to superclass Monadofilosa. A new order Tremulida was assigned to subphylum Filosa, Cercobodo became obsolete:[15]

In Cavalier-Smith and Karpov, 2012, superclass Monadofilosa was ranked as infraphylum. A new superclass called Ventrifilosa was used for the classes Thecofilosea and Imbricatea. A new class Skiomonadea was named for the only order Tremulida:[16]

In Yabuki et al., 2012, order Heliomonadida was transferred to class Endohelea and a new order called Axomonadida was added to class Granofilosea:[17]

In Cavalier-Smith and Oates, 2012, a new superclass Eoglissa was named to contain the classes Metromonadea and Sarcomonadea in the infraphylum Monadofilosa. Family Katabiidae was assigned class Sarcomonadea incertae sedis:[18]

In Cavalier-Smith and Chao, 2012, four new subclasses, three new superorders and four new orders were named. New subclass Placoperla was assigned to class Imbricatea and contained the new superorders Perlatia and Placofila. Superorder Perlatia included the order Spongomonadida and the new order Perlofilida. Superorder Placofila contained the orders Rotosphaerida and Thaumatomonadida. New subclass Placonuda was also assigned to class Imbricatea contained the order Euglyphida and the new superorder Nudisarca which included the order Marimonadida and the incertae sedis family Nudifilidae. New subclass Eothecia was assigned to class Thecofilosea and contained the orders Ventricleftida, Ebriida, Cryomonadida, Hemimastigida and new order Matazida. New subclass Tectosia was also assigned to class Thecofilosea and contained the infraclass Phaeodaria which was previously subclass Phaeodaria and order Tectofilosida. New orders Eodarida and Opaloconchida were assigned to infraclass Phaeodaria:[19]

In Adl et al., 2012:[20]

In Cavalier-Smith, 2013, order Rotosphaerida was treated as class Cristidiscoidea instead and was placed in its own new subphylum Paramycia in phylum Choanozoa:[21]

In Cavalier-Smith and Scoble, 2013, the incertae sedis genus Clautriavia was assigned to its own incertae sedis family Clautriaviidae:[22]

In Scoble and Cavalier-Smith, 2014, a new order Variglissida that included the families Nudifilidae and Clautriaviidae was named and placed in superorder Nudisarca. A new order Discomonadida that contained the only family Discomonadida and genus Discomonas was placed superclass Ventrifilosa incertae sedis. A new order Zoelucasida was placed in superorder Placofila. Order Pseudosporida was transferred to class Sarcomonadea:[23]

In Ruggiero et al., 2015, a generally simplified scheme was presented with also some taxa not listed:[24]

In Cavalier-Smith, 2018, a new order Minorisida was added to class Chlorarachnea. Order Axomonadida was used under class Granofilosea. Superclass Eoglissa was used for the class Metromonadea and new class Helkesea which included the order Ventricleftida and new order Helkesida that also included the superfamily Sainouroidea. Superclass Ventrifilosa was used for the classes Sarcomonadea, Imbricatea and Thecofilosea. Order Zoelucasida was transferred to subclass Placonuda and order Discocelida was transferred to subclass Placoperla. Order Perlofilida was still used but listed as only probably in subclass Placoperla. Subphylum Endomyxa was transferred to phylum Retaria:[25]

In Cavalier-Smith et al., 2018, order Aquavolonida was added to class Skiomonadea. Order Ventricleftida was transferred to a new subclass Ventricleftia in class Thecofilosea. A new order Paracercomonadida was assigned to a new subclass Paracercomonadia in class Sarcomonadea. Order Pansomonadida was ranked as suborder Pansomonadina and assigned to order Glissomonadida. Order Pseudosporida was assigned infrakingdom Rhizaria incertae sedis. A new subclass Pediglissa was assigned to class Sarcomonadea and contained the orders Cercomonadida and Glissomonadida. Superorder Nudisarca was used for the orders Variglissida and Marimonadida in subclass Placonuda. A new superorder Euglyphia was assigned to subclass Placonuda and contained the orders Euglyphida and Zoelucasida. A new superorder Discomonada was assigned to subclass Placonuda and contained the order Discomonadida. Superorder Placofila was used for the orders Thaumatomonadida and Discocelida in subclass Placoperla. Superorder Perlatia was used for the orders Spongomonadida and Perlofilida. A new subclass Krakenia was named to include a new order Krakenida in the class Imbricatea:[26]

In Adl et al., 2019:[27]

In Cavalier-Smith, 2022, order Axomonadida was assigned to class Endohelea and order Heliomonadida was assigned to class Granofilosea:[28]

References

  1. 1 2 Cavalier-Smith, T. (1998-08-01). "A revised six-kingdom system of life". Biological Reviews of the Cambridge Philosophical Society. 73 (3): 203–266. doi:10.1017/S0006323198005167.
  2. 1 2 Adl, Sina M.; Simpson, Alastair G. B.; Farmer, Mark A.; Andersen, Robert A.; Anderson, O. Roger; Barta, John R.; Bowser, Samuel S.; Brugerolle, Guy; Fensome, Robert A.; Fredericq, Suzanne; James, Timothy Y.; Karpov, Sergei; Kugrens, Paul; Krug, John; Lane, Christopher E. (2005). "The New Higher Level Classification of Eukaryotes with Emphasis on the Taxonomy of Protists". Journal of Eukaryotic Microbiology. 52 (5): 399–451. doi:10.1111/j.1550-7408.2005.00053.x. ISSN 1550-7408.
  3. ↑ Cavalier-Smith, T. (1997-04-01). "Amoeboflagellates and mitochondrial cristae in eukaryote evolution: megasystematics of the new protozoan subkingdoms eozoa and neozoa". Archiv für Protistenkunde. 147 (3): 237–258. doi:10.1016/S0003-9365(97)80051-6. ISSN 0003-9365.
  4. ↑ Cavalier-Smith, Thomas (1993). "The Protozoan Phylum Opalozoa". Journal of Eukaryotic Microbiology. 40 (5): 609–615. doi:10.1111/j.1550-7408.1993.tb06117.x. ISSN 1550-7408.
  5. ↑ Leadbeater, Barry S. C.; Green, John C., eds. (2000-06-08). Flagellates. CRC Press. doi:10.1201/9781482268225. ISBN 978-0-429-18213-6.
  6. 1 2 Bass, David; Chao, Ema E. -Y.; Nikolaev, Sergey; Yabuki, Akinori; Ishida, Ken-ichiro; Berney, Cédric; Pakzad, Ursula; Wylezich, Claudia; Cavalier-Smith, Thomas (2009-02-01). "Phylogeny of Novel Naked Filose and Reticulose Cercozoa: Granofilosea cl. n. and Proteomyxidea Revised". Protist. 160 (1): 75–109. doi:10.1016/j.protis.2008.07.002. ISSN 1434-4610.
  7. 1 2 Cavalier-Smith, Thomas; Chao, Ema E. -Y. (2003-10-01). "Phylogeny and Classification of Phylum Cercozoa (Protozoa)". Protist. 154 (3): 341–358. doi:10.1078/143446103322454112. ISSN 1434-4610.
  8. ↑ Cavalier-Smith, T (2002). "The phagotrophic origin of eukaryotes and phylogenetic classification of Protozoa". International Journal of Systematic and Evolutionary Microbiology. 52 (2): 297–354. doi:10.1099/00207713-52-2-297. ISSN 1466-5034.
  9. ↑ Bass, David; Cavalier-Smith, Thomas (2004). "Phylum-specific environmental DNA analysis reveals remarkably high global biodiversity of Cercozoa (Protozoa)". International Journal of Systematic and Evolutionary Microbiology. 54 (6): 2393–2404. doi:10.1099/ijs.0.63229-0. ISSN 1466-5034.
  10. ↑ Bass, David; Moreira, David; López-García, Purificación; Polet, Stephane; Chao, Ema E.; Heyden, Sophie von der; Pawlowski, Jan; Cavalier-Smith, Thomas (2005-08-18). "Polyubiquitin Insertions and the Phylogeny of Cercozoa and Rhizaria". Protist. 156 (2): 149–161. doi:10.1016/j.protis.2005.03.001. ISSN 1434-4610.
  11. ↑ Brodie, Juliet; Lewis, Jane, eds. (2007-11-26). Unravelling the algae: the past, present, and future of algal systematics (0 ed.). CRC Press. doi:10.1201/9780849379901. ISBN 978-0-429-12614-7.
  12. ↑ Cavalier-Smith, Thomas; Lewis, Rhodri; Chao, Ema E.; Oates, Brian; Bass, David (2008-10-21). "Morphology and Phylogeny of Sainouron acronematica sp. n. and the Ultrastructural Unity of Cercozoa". Protist. 159 (4): 591–620. doi:10.1016/j.protis.2008.04.002. ISSN 1434-4610.
  13. ↑ Howe, Alexis T.; Bass, David; Vickerman, Keith; Chao, Ema E.; Cavalier-Smith, Thomas (2009-05-01). "Phylogeny, Taxonomy, and Astounding Genetic Diversity of Glissomonadida ord. nov., The Dominant Gliding Zooflagellates in Soil (Protozoa: Cercozoa)". Protist. 160 (2): 159–189. doi:10.1016/j.protis.2008.11.007. ISSN 1434-4610.
  14. ↑ Cavalier-Smith, Thomas; Lewis, Rhodri; Chao, Ema E.; Oates, Brian; Bass, David (2009-08-01). "Helkesimastix marina n. sp. (Cercozoa: Sainouroidea superfam. n.) a Gliding Zooflagellate of Novel Ultrastructure and Unusual Ciliary Behaviour". Protist. 160 (3): 452–479. doi:10.1016/j.protis.2009.03.003. ISSN 1434-4610.
  15. ↑ Howe, Alexis T.; Bass, David; Scoble, Josephine M.; Lewis, Rhodri; Vickerman, Keith; Arndt, Hartmut; Cavalier-Smith, Thomas (2011-04-01). "Novel Cultured Protists Identify Deep-branching Environmental DNA Clades of Cercozoa: New Genera Tremula, Micrometopion, Minimassisteria, Nudifila, Peregrinia". Protist. 162 (2): 332–372. doi:10.1016/j.protis.2010.10.002. ISSN 1434-4610.
  16. ↑ Cavalier-Smith, Thomas; Karpov, Sergey A. (2012-01-01). "Paracercomonas Kinetid Ultrastructure, Origins of the Body Plan of Cercomonadida, and Cytoskeleton Evolution in Cercozoa". Protist. 163 (1): 47–75. doi:10.1016/j.protis.2011.06.004. ISSN 1434-4610.
  17. ↑ Yabuki, Akinori; Chao, Ema E.; Ishida, Ken-Ichiro; Cavalier-Smith, Thomas (2012-05-01). "Microheliella maris (Microhelida ord. n.), an Ultrastructurally Highly Distinctive New Axopodial Protist Species and Genus, and the Unity of Phylum Heliozoa". Protist. 163 (3): 356–388. doi:10.1016/j.protis.2011.10.001. ISSN 1434-4610.
  18. ↑ Cavalier-Smith, Thomas; Oates, Brian (2012-03-01). "Ultrastructure of Allapsa vibrans and the Body Plan of Glissomonadida (Cercozoa)". Protist. 163 (2): 165–187. doi:10.1016/j.protis.2011.10.006. ISSN 1434-4610.
  19. ↑ Cavalier-Smith, Thomas; Chao, Ema E. (2012-07-01). "Oxnerella micra sp. n. (Oxnerellidae fam. n.), a Tiny Naked Centrohelid, and the Diversity and Evolution of Heliozoa". Protist. 163 (4): 574–601. doi:10.1016/j.protis.2011.12.005. ISSN 1434-4610.
  20. ↑ Adl, Sina M.; Simpson, Alastair G. B.; Lane, Christopher E.; Lukeš, Julius; Bass, David; Bowser, Samuel S.; Brown, Matthew W.; Burki, Fabien; Dunthorn, Micah; Hampl, Vladimir; Heiss, Aaron; Hoppenrath, Mona; Lara, Enrique; le Gall, Line; Lynn, Denis H. (2012-09-28). "The Revised Classification of Eukaryotes". Journal of Eukaryotic Microbiology. 59 (5): 429–514. doi:10.1111/j.1550-7408.2012.00644.x. ISSN 1066-5234. PMC 3483872. PMID 23020233.
  21. ↑ Cavalier-Smith, Thomas (2013-05-01). "Early evolution of eukaryote feeding modes, cell structural diversity, and classification of the protozoan phyla Loukozoa, Sulcozoa, and Choanozoa". European Journal of Protistology. 49 (2): 115–178. doi:10.1016/j.ejop.2012.06.001. ISSN 0932-4739.
  22. ↑ Cavalier-Smith, Thomas; Scoble, Josephine Margaret (2013-08-01). "Phylogeny of Heterokonta: Incisomonas marina, a uniciliate gliding opalozoan related to Solenicola (Nanomonadea), and evidence that Actinophryida evolved from raphidophytes". European Journal of Protistology. 49 (3): 328–353. doi:10.1016/j.ejop.2012.09.002. ISSN 0932-4739.
  23. ↑ Scoble, Josephine M.; Cavalier-Smith, Thomas (2014-06-01). "Scale evolution, sequence phylogeny, and taxonomy of thaumatomonad Cercozoa: 11 new species and new genera Scutellomonas, Cowlomonas, Thaumatospina and Ovaloplaca". European Journal of Protistology. 50 (3): 270–313. doi:10.1016/j.ejop.2013.12.005. ISSN 0932-4739.
  24. ↑ 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. (2015-06-11). "Correction: A Higher Level Classification of All Living Organisms". PLOS ONE. 10 (6) e0130114. doi:10.1371/journal.pone.0130114. ISSN 1932-6203. PMC 5159126. PMID 26068874.
  25. ↑ Cavalier-Smith, Thomas (2018-01-01). "Kingdom Chromista and its eight phyla: a new synthesis emphasising periplastid protein targeting, cytoskeletal and periplastid evolution, and ancient divergences". Protoplasma. 255 (1): 297–357. doi:10.1007/s00709-017-1147-3. ISSN 1615-6102. PMC 5756292. PMID 28875267.
  26. ↑ Cavalier-Smith, Thomas; Chao, Ema E.; Lewis, Rhodri (2018-09-01). "Multigene phylogeny and cell evolution of chromist infrakingdom Rhizaria: contrasting cell organisation of sister phyla Cercozoa and Retaria". Protoplasma. 255 (5): 1517–1574. doi:10.1007/s00709-018-1241-1. ISSN 1615-6102. PMC 6133090. PMID 29666938.
  27. ↑ Adl, Sina M.; Bass, David; Lane, Christopher E.; Lukeš, Julius; Schoch, Conrad L.; Smirnov, Alexey; Agatha, Sabine; Berney, Cedric; Brown, Matthew W.; Burki, Fabien; Cárdenas, Paco; Čepička, Ivan; Chistyakova, Lyudmila; del Campo, Javier; Dunthorn, Micah (2019). "Revisions to the Classification, Nomenclature, and Diversity of Eukaryotes". Journal of Eukaryotic Microbiology. 66 (1): 4–119. doi:10.1111/jeu.12691. ISSN 1550-7408. PMC 6492006. PMID 30257078.
  28. ↑ Cavalier-Smith, Thomas (2021-12-23). "Ciliary transition zone evolution and the root of the eukaryote tree: implications for opisthokont origin and classification of kingdoms Protozoa, Plantae, and Fungi". Protoplasma. 259 (3): 487–593. doi:10.1007/s00709-021-01665-7. ISSN 0033-183X. PMC 9010356. PMID 34940909.
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