Axodine

Axodines
	An actinophryid
Scientific classification
Domain:	Eukaryota
Clade:	Diaphoretickes
Clade:	SAR
Clade:	Stramenopiles
Phylum:	Gyrista
Subphylum:	Ochrophytina
Class:	Dictyochophyceae
Subclass:	Pedinellia; Cavalier-Smith 1986 stat. nov. 2017[1]
Genera
	Dictyochales/Silicoflagellates Dictyocha; ; Rhizochromulinales Rhizochromulina; ; Actinodines; Pedinellales Pedinella; Apedinella; Pseudopedinella; Mesopedinella; Parapedinella; Actinomonas; Pteridomonas; Ciliophrys; ; Actinophryida; Actinophrys; Actinosphaerium; Florenciellales Pseudochattonella; ;
Synonyms
	Actinochrysea Cavalier-Smith 1995[2]; Axodines Patterson, 1994[3];

The axodines are a group of unicellular stramenopiles that includes silicoflagellate and rhizochromulinid algae, actinomonad heterotrophic flagellates and actinophryid heliozoa. Alternative classifications treat the dictyochophytes as heterokont algae, or as Chrysophyceae.[4] Other overlapping taxonomic concepts include the Actinochrysophyceae, Actinochrysea or Dictyochophyceae sensu lato. The grouping was proposed on the basis of ultrastructural similarities, and is consistent with subsequent molecular comparisons.[5]

The Axodine grouping was unusual in breaking with the traditions of botanical and protozoological taxonomy to include the actinophryid heliozoa as part of the lineage that also contained the pedinellid algae along with colorless relatives such as Actinomonas, Pteridomonas, and Ciliophrys; the axodines further included the silicoflagellates, and Rhizochromulinales.[3] This followed a growing consensus that the actinophryid heliozoa were not related to other types of heliozoa.[6][7][8]

Characteristics

The name points to a character that is deemed to be synapomorphic for the group: that is the microtubular arrays that extend from the surface of the nucleus. Many flagellated forms have a single emergent flagellum, that lacks the root structure found in related chrysophytes.

Siliceous skeleton of the dictyochid flagellate, Dictyocha fibula
Actinophrys sol, living heliozoon
Actinomonas - heterotrophic flagellate with single flagellum and stiff arms
Ciliophrys infusionum, living flagellate

Classification and history

Diagram by Mikrjukov and Patterson to show the relationships of, and within, axodines, together with the proposed synapomorphies of the clades.

The traditional botanical treatment of the group follows

Order Dictyochales

The most notable group is the silicoflagellates, marine plankton that form siliceous skeletons and are well known as fossils.

Order Pedinellales

Most other axodines form a group variously called the pedinellids.

Order Rhizochromulinales

In addition to the silicoflagellates and actinodines, the marine amoeboid Rhizochromulina marina is included here based on the structure of its zoospores. It is considered closer to the latter group than the former.

References

Cavalier-Smith, Thomas (2017). "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. PMC 5756292. PMID 28875267.
Cavalier-Smith, T.; Chao, E.E.; Allsopp, M.T.E.P. (1995). "Ribosomal RNA evidence for chloroplast loss within Heterokonta: pedinellid relationships and a revised classification of ochristan algae". Arch. Protistenkd. 145 (3–4): 209–220. doi:10.1016/S0003-9365(11)80316-7.
Mikrjukov KA, Patterson DJ (1 February 2001). "Taxonomy and phylogeny of Heliozoa. III. Actinophryids" (PDF). Acta Protozoologica. 40 (1): 3–26.
Sandgren, C.D.; Smol, J.P.; Kristiansen, J. (1995). Chrysophyte Algae: Ecology, Phylogeny and Development. Cambridge University Press. p. 49. ISBN 9780521462600. Retrieved 2014-12-12.
Nikolaev, S.I., Berney, C., Fahrni, J., Bolivar, I.,Polet, S., Mylnikov, A.P., Aleshin, V.V., Petrov, N. B., Pawlowski, J. 2004. The twilight of Heliozoa and rise of Rhizaria, an emerging supergroup of amoeboid eukaryotes. Proceedings of the National Academy of Sciences 101 (21) 8066-8071; DOI: 10.1073/pnas.0308602101
Patterson, D. J. and Fenchel, T. 1985. Insights into the evolution of heliozoa (Protozoa, Sarcodina) as provided by ultrastructural studies on a new species of flagellate from the genus Pteridomonas. Biol. J. Linn. Soc. 34: 381-403
Patterson, D. J. 1986. The actinophryid heliozoa (Sarcodina, Actinopoda) as chromophytes. pp. 49 - 67 in Kristiansen, J. & Anderson, R. A. Chrysophytes: aspects and problems, Cambridge University Press, Cambridge & New York.
Smith, R. McK. & Patterson, D. J. 1986. Analyses of heliozoan inter-relationships: an example of the potentials and limitations of the ultrastructural approaches to the study of protistan phylogeny. Proc. R. Soc. Lond. 227:325 - 366.

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[8phyla-1] Cavalier-Smith, Thomas (2017). "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. PMC 5756292. PMID 28875267.

[2] Cavalier-Smith, T.; Chao, E.E.; Allsopp, M.T.E.P. (1995). "Ribosomal RNA evidence for chloroplast loss within Heterokonta: pedinellid relationships and a revised classification of ochristan algae". Arch. Protistenkd. 145 (3–4): 209–220. doi:10.1016/S0003-9365(11)80316-7.

[TaxoHeliozoa3-3] Mikrjukov KA, Patterson DJ (1 February 2001). "Taxonomy and phylogeny of Heliozoa. III. Actinophryids" (PDF). Acta Protozoologica. 40 (1): 3–26.

[google-4] Sandgren, C.D.; Smol, J.P.; Kristiansen, J. (1995). Chrysophyte Algae: Ecology, Phylogeny and Development. Cambridge University Press. p. 49. ISBN 9780521462600. Retrieved 2014-12-12.

[5] Nikolaev, S.I., Berney, C., Fahrni, J., Bolivar, I.,Polet, S., Mylnikov, A.P., Aleshin, V.V., Petrov, N. B., Pawlowski, J. 2004. The twilight of Heliozoa and rise of Rhizaria, an emerging supergroup of amoeboid eukaryotes. Proceedings of the National Academy of Sciences 101 (21) 8066-8071; DOI: 10.1073/pnas.0308602101

[6] Patterson, D. J. and Fenchel, T. 1985. Insights into the evolution of heliozoa (Protozoa, Sarcodina) as provided by ultrastructural studies on a new species of flagellate from the genus Pteridomonas. Biol. J. Linn. Soc. 34: 381-403

[7] Patterson, D. J. 1986. The actinophryid heliozoa (Sarcodina, Actinopoda) as chromophytes. pp. 49 - 67 in Kristiansen, J. & Anderson, R. A. Chrysophytes: aspects and problems, Cambridge University Press, Cambridge & New York.

[8] Smith, R. McK. & Patterson, D. J. 1986. Analyses of heliozoan inter-relationships: an example of the potentials and limitations of the ultrastructural approaches to the study of protistan phylogeny. Proc. R. Soc. Lond. 227:325 - 366.