Rhizammina algaeformis, from Brady (1884).

Belongs within: Foraminifera.

If a komokiacean turns up in a phylogeny, will anybody notice?
Published 13 July 2010

It’s possible that only one reader will care about this. But this is one minor detail that I came across while researching yesterday’s post.

Komokiacea are large deep-sea branching protists that are expected, but not conclusively demonstrated, to belong or be related to Foraminifera. Attempts to extract molecular data for komokiaceans have, to date, failed miserably (Lecroq et al. 2009). Or have they?

In a paper written by Gooday et al. (1997), my attention was caught by the offhand comment: “We include Rhizammina algaeformis Brady 1879 [another deep-sea branching foram] within the Komokiacea on the basis of its ‘softpart’ organization“. This relationship, it turns out, had originally been proposed by Cartwright et al. (1989). If this assignment is correct then komokiaceans have been appearing in molecular phylogenies for some years and nobody has been paying a blind bit of notice! Even more interestingly, Pawlowski et al. (2003) placed Rhizammina as sister to the xenophyophore Syringammina. Xenophyophores resemble komokiaceans both in being large branchers and in the sequestration of stercomata (faecal pellets) within their structure. A close relationship between the two groups would be quite credible. However, no other authors to date appear to have commented on Cartwright et al.‘s reclassification of Rhizammina so I have no idea whether it’s regarded as credible.


Systematics of Dendrophryida
    |    |--Halyphysema Bowerbank 1862LT64 (see below for synonymy)
    |    |    `--*H. tumanowiczii Bowerbank 1862 (see below for synonymy)LT64
    |    |--SchizamminidaeLT64
    |    |    |--Jullienella Schlumberger 1890LT64
    |    |    |    |--*J. foetida Schlumberger 1890LT64
    |    |    |    `--J. zealandicaPG98
    |    |    `--Schizammina Heron-Allen & Earland 1929 (see below for synonymy)LT64
    |    |         |--*S. labyrinthica Heron-Allen & Earland 1929LT64
    |    |         `--S. wedmoriensis (Chapman 1895) (see below for synonymy)LT64
    |    `--Diffusilinidae [Diffusilininae]M13
    |         |--Diffusilina Heron-Allen & Earland 1924LT64
    |         |    `--*D. humilis Heron-Allen & Earland 1924LT64
    |         |--Discobotellina Collins 1958LT64
    |         |    `--*D. biperforata Collins 1958LT64
    |         |--Kerionammina Moreman 1933LT64
    |         |    `--*K. favus Moreman 1933LT64
    |         |--Masonella Brady 1889 [=Armasonellum Rhumbler 1913]LT64
    |         |    `--*M. planulata Brady 1889 [=*Armasonellum planulatum]LT64
    |         |--Pseudowebbinella Shchedrina 1962LT64
    |         |    `--*P. goesi (Höglund 1947) [=Crithionina goesi]LT64
    |         |--Thuramminoides Plummer 1945LT64
    |         |    `--*T. sphaeroidalis Plummer 1945LT64
    |         |--Verrucina Goës 1896 [=Arverrucum Rhumbler 1913]LT64
    |         |    `--*V. rudis Goës 1896 [=*Arverrucum rude]LT64
    |         `--Weikkoella Summerson 1958LT64
    |              `--*W. sphaerica Summerson 1958LT64
    `--Dendrophryoidea [Komokiacea]M13
         |  i. s.: Clados Schroeder, Medioli & Scott 1989M13
         |--Dryorhizopsis Henbest 1963HW93 [DryorhizopsidaeM13]
         |--NotodendrodesPH03 [NotodendrodidaeM13]
         |    `--N. antarctikosPH03
         |    |--Arborammina hilaryaeRR96
         |    `--Luffammina Kamenskaya, Bagirov & Simdianov 2002M13
         |--Normanina Cushman 1928TH77 [NormaninidaeM13]
         |    |--*N. conferta (Norman 1878) [=Haliphysema confertum]TH77
         |    |--N. saidovae Tendal & Hessler 1977TH77
         |    `--N. tylota Tendal & Hessler 1977TH77
         |    |--Baculella Tendal & Hessler 1977TH77
         |    |    |--*B. globofera Tendal & Hessler 1977TH77
         |    |    `--B. hirsuta Tendal & Hessler 1977TH77
         |    `--Edgertonia Tendal & Hessler 1977TH77
         |         |--*E. tolerans Tendal & Hessler 1977TH77
         |         `--E. argillispherula Tendal & Hessler 1977TH77
         |    |--Spiculidendron Rützler & Richardson 1996RR96 [SpiculidendroninaeM13]
         |    |    `--*S. corallicolum Rützler & Richardson 1996RR96
         |    `--DendrophryinaeM13
         |         |--Dendronina Heron-Allen & Earland 1922LT64
         |         |    `--*D. arborescens Heron-Allen & Earland 1922LT64
         |         |--Nubeculariella Averintsev 1911LT64
         |         |    `--*N. birulai Averintsev 1911LT64
         |         `--Dendrophrya Wright 1861 [=Ardendrophyrum Rhumbler 1913, Dendrophyra (l. c.)]LT64
         |              `--*D. erecta Wright 1861LT64
         `--Komokiidae [Arrhizamnia, Rhizamminidae, Rhizammininae, Testulosiphoninae]M13
              |--Calos Schroeder, Medioli & Scott 1989M13
              |--Cactos Schroeder, Medioli & Scott 1989M13
              |--Cerebrum Schroeder, Medioli & Scott 1989M13
              |--Crambis Schroeder, Medioli & Scott 1989M13
              |--‘Reticulum’ Schroeder, Medioli & Scott 1989 non de Stefani 1879M13
              |--Tuber Schroeder, Medioli & Scott 1989 non Wigg. 1780 (ICBN)M13
              |--Skeletonia Gooday, Kamenskaya & Cedhagen 2007M13
              |--Komokia Tendal & Hessler 1977TH77
              |    `--*K. multiramosa Tendal & Hessler 1977TH77
              |--Septuma Tendal & Hessler 1977TH77
              |    `--*S. ocotillo Tendal & Hessler 1977TH77
              |--Ipoa Tendal & Hessler 1977TH77
              |    `--*I. fragila Tendal & Hessler 1977TH77
              |--Marsipella Norman 1878 [=Armarsipellum Rhumbler 1913]LT64
              |    |--*M. elongata Norman 1878 [=*Armarsipellum elongatum]LT64
              |    `--M. aggregata Moreman 1933C61
              |--Lana Tendal & Hessler 1977TH77
              |    |--*L. neglecta Tendal & Hessler 1977TH77
              |    `--L. reticulata Tendal & Hessler 1977TH77
              `--Rhizammina Brady 1879M13, LT64 (see below for synonymy)
                   |--*R. algaeformis Brady 1879 [=*Arrhizammum algaeforme]LT64
                   |--R. globigerinifera Hofker 1930 [=*Testulorhiza globigerinifera]LT64
                   `--R. indivisa Brady 1884 [=*Testulosiphon indivisum]LT64
Nomina nuda: Normanina elongata Saidova 1968TH77
             Normanina fruticosa Saidova 1968TH77
             Normanina ultraabyssalica Saidova 1968TH77

Halyphysema Bowerbank 1862LT64 [=Arhaliphysemum Rhumbler 1913LT64, Gastrophysema Haeckel 1877LT64, Haliphysema Haeckel 1877LT64; HalyphysematidaeM13]

*Halyphysema tumanowiczii Bowerbank 1862 [=*Arhaliphysemum tumanowiczii, *Haliphysema tumanowiczii, Squamulina scopula Carter 1877, *Gastrophysema scopula]LT64

Rhizammina Brady 1879M13, LT64 [=Arrhizammum Rhumbler 1913LT64; incl. Testulorhiza Avnimelech 1952LT64, Testulosiphon Avnimelech 1952LT64]

Schizammina Heron-Allen & Earland 1929 [incl. Arpsammosiphoum Rhumbler 1913, Psammosiphon Rhumbler 1911 non Vine 1882]LT64

Schizammina wedmoriensis (Chapman 1895) [=Nodosinella wedmoriensis, *Arpsammosiphoum wedmoriense, *Psammosiphon wedmoriense]LT64

*Type species of generic name indicated


Cartwright, N. G., A. J. Gooday & A. R. Jones. 1989. The morphology, internal organization, and taxonomic position of Rhizammina algaeformis Brady, a large, agglutinated, deep-sea foraminifer. Journal of Foraminiferal Research 19 (2): 115–125.

[C61] Crespin, I. 1961. Upper Devonian Foraminifera from Western Australia. Palaeontology 3 (4): 397–409.

Gooday, A. J., R. Shires & A. R. Jones. 1997. Large, deep-sea agglutinated Foraminifera; two differing kinds of organization and their possible ecological significance. Journal of Foraminiferal Research 27 (4): 278–291.

[HW93] Hart, M. B., & C. L. Williams. 1993. Protozoa. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 43–70. Chapman & Hall: London.

Lecroq, B., A. J. Gooday, T. Cedhagen, A. Sabbatini & J. Pawlowski. 2009. Molecular analyses reveal high levels of eukaryotic richness associated with enigmatic deep-sea protists (Komokiacea). Marine Biodiversity 39: 45–55.

[LT64] Loeblich, A. R., Jr & H. Tappan. 1964. Sarcodina: chiefly “thecamoebians” and Foraminiferida. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt C. Protista 2 vol. 1. The Geological Society of America and The University of Kansas Press.

[M13] Mikhalevich, V. I. 2013. New insight into the systematics and evolution of the Foraminifera. Micropaleontology 59 (6): 493–527.

[PH03] Pawlowski, J., M. Holzmann, J. Fahrni & S. L. Richardson. 2003. Small subunit ribosomal DNA suggests that the xenophyophorean Syringammina corbicula is a foraminiferan. Journal of Eukaryotic Microbiology 50: 483–487.

[PG98] Probert, P. K., & S. L. Grove. 1998. Macrobenthic assemblages of the continental shelf and upper slope off the west coast of South Island, New Zealand. Journal of the Royal Society of New Zealand 28: 259–280.

[RR96] Rützler, K., & S. Richardson. 1996. The Caribbean spicule tree: a sponge-imitating foraminifer (Astrorhizidae). Bulletin de l’Institut Royal des Sciences Naturelles de Belgique 66 (Suppl.): 143–151.

[TH77] Tendal, O. S., & R. R. Hessler. 1977. An introduction to the biology and systematics of Komokiacea (Textulariina, Foraminiferida). Galathea Report 14: 165–194.

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