Saccamminidae

Pelosina variabilis, copyright Jan Pawlowski.

Belongs within: Globothalamea.

The Saccamminidae are a group of Foraminifera with agglutinated tests known from the Ordovician to the present (Loeblich & Tappan 1964); it is likely that this represents a polyphyletic assemblage of relatively simple-bodied forams.

A small bag of grains
Published 9 March 2010
Tests of the agglutinating foram Saccammina sphaerica. Scale bar = 500 μm. These are the lectotype and paralectotype of this species held by the British Natural History Museum.

Foraminifera are one of the best-known of protist groups and may make up more than half of the benthic biomass in some marine habitats, particularly in the deep sea (Gooday et al. 2001). Forams are amoeboids with filose pseudopodia that branch and rejoin each other to form a net for the collection of food particles. The great majority of forams are marine, and most (but not all) forams produce some sort of protective test or shell with the filopodia extending from openings or pores in the test. Because of this test, forams are one of the few protist groups with an extensive fossil record. Indeed, their use in biostratigraphic studies (and leading surveyors to small treats such as oil deposits) has lead to forams being better known from a palaeontological than a Recent perspective and the structure and morphology of the test has long been a major factor in distinguishing and classifying forams.

Forams may secrete their own tests (usually chitinous or calcareous) or they may construct a test from sand grains and/or other foreign particles (these are known as agglutinating forams). A distinction is also commonly made between monothalamous forms, in which the test is not divided into chambers (at least, not by complete septa), and polythalamous forms, in which the test is divided by septa into a number of chambers. However, molecular phylogenetic studies of recent forams have shown that monothalamous forams are paraphyletic while polythalamous forms are potentially polyphyletic (Flakowski et al. 2005). Also, while monothalamous taxa may produce chitinous or agglutinated tests, the type of test produced does not appear to correspond with phylogenetic position (Pawlowski et al. 2002).

Despite this, foram researchers continue to use the old test-based classification for the simple reason that no-one has yet come up with a better alternative (and doing so would not be easy). The Saccamminidae as generally recognised are a large family of agglutinating forams with generally a single chamber (or sometimes a bunch of similar chambers loosely attached to each other), usually with a single aperture. They may be globular or more elongate in shape. Some saccamminids are quite catholic in their choice of building materials but others may be more fussy. Perhaps the ultimate in fussiness is expressed by Technitella thompsoni which builds its test with nothing but the ambulacral plates of brittle stars (Cushman 1940).

The ‘silver saccamminid’, an as-yet unidentified species that has appeared in a number of phylogenetic studies. Photo by Jan Pawlowski.

As with other monothalamous groups, molecular phylogenetic studies have indicated that “saccamminids” are polyphyletic with representative species scattered in various positions among the basal part of the foram tree (Cedhagen et al. 2009; Gooday & Pawlowski 2004). Despite a fossil record extending back to the Cambrian (with putative ‘saccamminids’ at least as far back as the Silurian) and a significant abundance in the modern marine benthos, agglutinating forams are not as well-studied as calcareous taxa and monothalamous forms are particularly poorly so. For a start, they are often extremely small. Ecological studies have found the majority of ‘saccamminid’ specimens to be much less than 100 μm in diameter (Gooday et al. 2001) though Pilulina jeffreysii reaches more than 4 mm (Cedhagen et al. 2009). Also problematic, particularly for palaeontological studies, is that agglutinating foram tests are often difficult to distinguish from their surrounding environment because they are, after all, made from their surrounding environment. Perhaps the best demonstration of this issue is that the Stannomidae, despite being the largest of all forams and reaching over a foot in size, have no recognised fossil history at all.

Three random foram genera
Published 12 July 2010

As explained above, the ‘Saccamminidae’ are undoubtedly a polyphyletic assemblage of forams of very simple morphology. In the influential, and outdated, classification of Cushman (1940), ‘saccamminids’ were divided between four subfamilies for which odds are that each of those subfamilies are as polyphyletic as the whole. Let’s take a look at the members of one of those subfamilies and see where they are now.

The subfamily Pelosininae, as recognised by Cushman (1940), included the genera Pelosina, Technitella and Pilulina. The distinguishing characteristics of this subfamily were that its members had free, unattached tests with a single chamber, at least one aperture and walls composed of fine particles. All three also live in the deep sea and include relatively large species for forams (up to 60 mm in height in Pelosina). In the classification of Kaminksi (2004), none of these genera were closely associated. In the case of Pelosina, Cushman was not even correct about the few defining features of the subfamily because this genus does live attached to the sediment by root-like structures (Rützler & Richardson 1996). Pelosina species are one of a number of tree-like forams that form a significant component of the deep-sea benthic community.

Technitella legumen, from here.

Technitella, in contrast, is an elongate, somewhat sausage-like form. The name of the genus (“little workman”) refers to its elegantly constructed test, constructed from carefully selected materials. Heron-Allen & Earland (1909) described one species, T. thompsoni, which uses nothing but brittle star plates while T. legumen prefers sponge spicules, arranged in two layers with the spicules in each layer at right angles to each other to strengthen the test. Heron-Allen and Earland mused that “Probably we should be considered as imposing too weighty a postulate upon the members of the Club if we ventured to suggest that these rudimentary organisms were gifted with any aesthetic sense… it would appear that this “primordial, protoplasmic, atomic globule” is by no means so elementary an organism as naturalists are inclined to believe“.

Lectotype and paralectotype of Pilulina jeffreysii, photographed by Andrew Henderson.

Finally, Pilulina constructs a globular test of felted sponge spicules with an elongate aperture like the mouth on a stick-figure’s face. Of the three genera, only Pelosina and Pilulina have appeared in molecular phylogenetic analyses and the two do not appear to be associated, sitting instead in separate parts of the saccamminid cloud (e.g., Lecroq et al., 2009). Mikhalevich & Voronova (1999) argued that Pelosina is in fact a xenophyophore and placed it in the order Stannomida with the genera Stannoma and Stannophyllum. This was based on the supposed presence of linellae, a structure only otherwise found in stannomids. No molecular analysis has indicated an association between Pelosina and other xenophyophores. However, no other stannomid has appeared in these analyses, so just because Pelosina is not directly related to xenophyophores may not necessarily mean that it is not directly related to stannomids.

Systematics of Saccamminidae

Characters (from Loeblich & Tappan 1964): Test free or attached, subglobular, or in groups; aperture absent, single, or multiple.

<==Saccamminidae [Kyphamminidae, Pelosininae, Pilulinida, Pilulinidae, Pilulinina, Protocystidae, Saccamminae, Saccamminina]
    |--SaccammininaeM13
    |    |--Saccammina Sars in Carpenter 1869 (see below for synonymy)LT64
    |    |    |--*S. sphaerica Brady 1871 [=*Arsaccammum sphaericum]LT64
    |    |    |--S. lathramiBL79
    |    |    `--‘Reophax’ placenta Grzybowski 1897 [=*Placentammina placenta]LT64
    |    `--Lavella Nestell & Tolmacheva 2004M13
    |--OvammininaeM13
    |--Pelosina Brady 1879LT64 (see below for synonymy)
    |    |--*P. variabilis Brady 1879 [=*Arpelosum variabile]LT64
    |    |--P. arborescensG96
    |    |--P. bicaudata (Parr 1950) [=*Pelosinella bicaudata]LT64
    |    |--P. caudata (Montanaro Gallitelli 1955) [=Saccammina caudata, *Caudammina caudata]LT64
    |    |--P. distoma Millett 1904 [=*Millettina distoma]LT64
    |    |--P. rotundataC40
    |    `--P. sphaeriloculum Höglund 1947 [=P. variabilis var. sphaeriloculum, *Globosiphon sphaeriloculum]LT64
    `--Pilulina Carpenter 1870LT64 [=Arpilulum Rhumbler 1913LT64; PilulininaeM13]
         `--*P. jeffreysii Carpenter 1870 [=*Arpilulum jeffreysii]LT64
Saccamminidae incertae sedis:
  Pelosphaera Heron-Allen & Earland 1932LT64
    `--*P. cornuta Heron-Allen & Earland 1932LT64
  Saccamminoides Geroch 1955LT64
    `--*S. carpathicus Geroch 1955LT64
  Stomasphaera Mound 1961LT64
    `--*S. brassfieldensis Mound 1961LT64
  Lagenammina Rhumbler 1911 [=Arlagenammum Rhumbler 1913]LT64
    |--*L. laguncula Rhumbler 1911 [=*Arlagenammum laguncula]LT64
    `--L. compressa (Cushman & McCulloch 1939) [=Proteonina compressa]H03
  Technitella Norman 1878 (see below for synonymy)LT64
    |--*T. legumen Norman 1878 [=*Artechnitum legumen]LT64
    `--T. richardi (de Folin 1887) [=*Dioxeia richardi]LT64
  Brachysiphon Chapman 1906 (see below for synonymy)LT64
    |--*B. corbuliformis Chapman 1906 [=*Arbrachysiphum corbuliforme]LT64
    |--B. australe (Crespin 1958) [=*Sacculinella australe]LT64
    `--B. rudis (Parr 1942) [=Hyperammina rudis, *Hyperamminita rudis]LT64
  Ordovicina Eisenack 1938 (see below for synonymy)LT64
    |--*O. oligostoma Eisenack 1938LT64
    |--O. bicuspidata (Dunn 1942) [=*Shidelerella bicuspidata]LT64
    |--O. citroniforma (Eisenack 1938) [=*Amphitremoida citroniforma, *Amphitremoidea citroniforma]LT64
    |--O. typa (Dunn 1942) [=*Croneisella typa]LT64
    `--O. williamsae (Dunn 1942) [=*Gastroammina williamsae]LT64

Brachysiphon Chapman 1906 [=Arbrachysiphum Rhumbler 1913; incl. Hyperamminita Crespin 1958, Sacculinella Crespin 1958]LT64

Ordovicina Eisenack 1938 [incl. Amphitremoida Eisenack 1938, Amphitremoidea Thalmann 1941, Croneisella Dunn 1942, Gastroammina Dunn 1942, Shidelerella Dunn 1942]LT64

Pelosina Brady 1879LT64 [=Arpelosum Rhumbler 1913LT64; incl. Caudammina Montanaro Gallitelli 1955LT64, Globosiphon Avnimelech 1952LT64, Millettina Avnimelech 1952LT64, Pelosinella Parr 1950LT64; CaudammininaeM13]

Saccammina Sars in Carpenter 1869 [=Arsaccammum Rhumbler 1913, Saccamina (l. c.); incl. Placentammina Majzon 1943]LT64

Technitella Norman 1878 [=Artechnitum Rhumbler 1913; incl. Dioxeia de Folin 1887, Hyperamminella de Folin 1887]LT64

*Type species of generic name indicated

References

[BL79] Basov, V. A., B. G. Lopatin, I. S. Gramberg, A. I. Danjushevskaya, V. Ya. Kaban’kov, V. M. Lazurkin & D. K. Patrunov. 1979. Lower Cretaceous lithostratigraphy near Galicia Bank. Initial Reports of the Deep Sea Drilling Project 47: 683–717.

Cedhagen, T., A. J. Gooday & J. Pawlowski. 2009. A new genus and two new species of saccamminid foraminiferans (Protista, Rhizaria) from the deep Southern Ocean. Zootaxa 2096: 9–22.

[C40] Cushman, J. A. 1940. Foraminifera: Their classification and economic use 3rd ed. Harvard University Press: Cambridge (Massachusetts).

Flakowski, J., I. Bolivar, J. Fahrni & J. Pawlowski. 2005. Actin phylogeny of Foraminifera. Journal of Foraminiferal Research 35 (2): 93–102.

[G96] Gooday, A. J. 1996. Xenophyophores (Protista), including two new species, from two abyssal sites in the northeast Atlantic Ocean. Journal of Foraminiferal Research 26 (3): 193–208.

Gooday, A. J., H. Kitazato, S. Hori & T. Toyofuku. 2001. Monothalamous soft-shelled Foraminifera at an abyssal site in the North Pacific: a preliminary report. Journal of Oceanography 57: 377–384.

Gooday, A. J., & J. Pawlowski. 2004. Conqueria laevis gen. and sp. nov., a new soft-walled, monothalamous foraminiferan from the deep Weddell Sea. Journal of the Marine Biological Association of the UK 84 (5): 919–924.

[H03] Hanagata, S. 2003. Miocene-Pliocene Foraminifera from the Niigata oil-fields region, northeastern Japan. Micropaleontology 49 (4): 293–340.

Heron-Allen, E., & A. Earland. 1909. On a new species of Technitella from the North Sea, with some observations upon selective power as exercised by certain species of arenaceous Foraminifera. Journal of the Quekett Microscopical Club, second series 10: 403–412.

Kaminski, M. A. 2004. The Year 2000 classification of the agglutinated Foraminifera. In: Bubík, M., & M. A. Kaminski (eds) Proceedings of the Sixth International Workshop on Agglutinated Foraminifera. Grzybowski Foundation Special Publication 8: 237–255.

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.

Mikhalevich, V. I., & M. N. Voronova. 1999. O sistematicheskom polozhenii roda Pelosina (Xenophyophoria, Protista, inc. sedis). Zoologicheskii Zhurnal 78 (2): 133–141.

Pawlowski, J., M. Holzmann, C. Berney, J. Fahrni, T. Cedhagen & S. S. Bowser. 2002. Phylogeny of allogromiid Foraminifera inferred from SSU rRNA gene sequences. Journal of Foraminiferal Research 32 (4): 334–343.

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.

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