Foraminifera

Foraminifera (/frmnfr/, Latin meaning hole

bearers, informally called "forams") are members of a
phylum or class of amoeboid protists characterized by
streaming granular ectoplasm that among other things is
used for catching food, and commonly by an external shell
or "test" made of various materials and constructed in
diverse forms. All but perhaps a very few are aquatic
and most are marine, the majority of which live on or
within the seaoor sediment (i.e., are benthic) while a
smaller variety are oaters in the water column at various depths (i.e., are planktonic). A few are known from
freshwater or brackish conditions and some soil species
have been identied through molecular analysis of small
subunit ribosomal DNA.[1][2]

in which they had been placed.

Although as yet unsupported by morphological correlates, molecular data strongly suggest that Foraminifera
are closely related to the Cercozoa and Radiolaria, both of
which also include amoeboids with complex shells; these
three groups make up the Rhizaria.[9] However, the exact
relationships of the forams to the other groups and to one
another are still not entirely clear.

2 Living foraminifera
Modern foraminifera are primarily marine organisms but
living individuals have been found in brackish, freshwater [13] and even terrestrial habitats.[2] The majority
of the species are benthic, and a further 40 morphospecies are planktonic.[14] This count may however represent only a fraction of actual diversity, since many
genetically discrepant species may be morphologically
indistinguishable.[15]

Foraminifera typically produce a test, or shell, which can

have either one or multiple chambers, some becoming
quite elaborate in structure.[3] These shells are commonly
made of calcium carbonate (CaCO
3) or agglutinated sediment particles. Over 10,000
species are recognized, both living (8,708) and fossil
(1,837).[4] They are usually less than 1 mm in size, but
some are much larger, the largest species reaching up to A number of forams have unicellular algae as
endosymbionts, from diverse lineages such as the
20 cm.[5]
green algae, red algae, golden algae, diatoms, and
dinoagellates.[14] Some forams are kleptoplastic,
retaining chloroplasts from ingested algae to conduct
1 Classication-taxonomy
photosynthesis.[16]

The taxonomic position of Foraminifera has varied since

their recognition as protozoa (protists) by Schultze in 3 Biology
1854,[6] there referred to as an order, Foraminiferida.
Loeblich and Tappan (1992) re-ranked Foraminifera as The foraminiferal cell is divided into granular endoplasm
a class[7] as it is now commonly regarded.
and transparent ectoplasm from which a pseudopodial
Foraminifera have typically been included in the net may emerge through a single opening or through
Protozoa,[8][9][10] or in the similar Protoctista or Protist many perforations in the test. Individual pseudopods
kingdom.[11][12] There is compelling evidence, based characteristically have small granules streaming in both
primarily on molecular phylogenetics, for their be- directions.[13] The pseudopods are used for locomotion,
longing to a major group within the Protozoa known anchoring, and in capturing food, which consists of small
as the Rhizaria.[8] Prior to the recognition of evo- organisms such as diatoms or bacteria.[14]
lutionary relationships among the members of the The foraminiferal life-cycle involves an alternation beRhizaria, Foraminifera were generally grouped with other tween haploid and diploid generations, although they are
Amoeboids as phylum Rhizopodea (or Sarcodina) in the mostly similar in form.[6][17] The haploid or gamont iniclass Granuloreticulosa.
tially has a single nucleus, and divides to produce nuRhizaria is problematic as it is often called a supergroup, rather than using an established taxonomic rank
such as phylum. Cavalier-Smith denes Rhizaria as an
infrakingdom within the Kingdom Protozoa.[8]

merous gametes, which typically have two agella. The

diploid or schizont is multinucleate, and after meiosis
fragments to produce new gamonts. Multiple rounds of
asexual reproduction between sexual generations is not
[13]
Some taxonomies put Foraminifera in a phylum of their uncommon in benthic forms.
own, putting them on par with the amoeboid Sarcodina Abundance of certain Foraminifera is sometimes used by
1

4 TESTS

researchers as an indicator of the completeness of vertical

mixing in certain seas such as the Celtic Sea.

Tests

The miliolid foraminiferan Quinqueloculina from the Belgian

part of the North Sea.

Foraminiferan tests (ventral view)

Thin section of a peneroplid foraminiferan from Holocene lagoonal sediment in Rice Bay, San Salvador Island, Bahamas.
Scale bar 100 micrometres.

Fossil nummulitid foraminiferans showing microspheric and

megalospheric individuals; Eocene of the United Arab Emirates;
scale in mm.

The form and composition of the test is the primary

means by which forams are identied and classied. Most
have calcareous tests, composed of calcium carbonate.[13]
In other forams the test may be composed of organic material, made from small pieces of sediment cemented together (agglutinated), and in one genus of silica. Openings in the test, including those that allow cytoplasm to
ow between chambers, are called apertures. The test
contains an organic matrix, which can sometimes be recovered from fossil samples.[18]

Ammonia beccarii, a benthic foram from the North Sea.

up the pyramids of Egypt is composed almost entirely

of nummulitic benthic Foraminifera.[20] Production estimates indicate that reef Foraminifera annually generate
approximately 43 million tons of calcium carbonate per
year and thus play an essential role in the production of
[21]
Tests are known as fossils as far back as the Cambrian reef carbonates.
period,[19] and many marine sediments are composed pri- Genetic studies have identied the naked amoeba
marily of them. For instance, the limestone that makes Reticulomyxa and the peculiar xenophyophores as

foraminiferans without tests. A few other amoeboids produce reticulose pseudopods, and were formerly classied with the forams as the Granuloreticulosa, but this is
no longer considered a natural group, and most are now
placed among the Cercozoa.[22]

ply of these fossil tests and the relatively high-precision

age-control models available for cores has produced an
exceptionally high-quality planktonic Foraminifera fossil
record dating back to the mid-Jurassic, and presents an
unparalleled record for scientists testing and documenting
the evolutionary process.[24] The exceptional quality of
the fossil record has allowed an impressively detailed picture of species inter-relationships to be developed on the
basis of fossils, in many cases subsequently validated independently through molecular genetic studies on extant
specimens[25] Larger benthic Foraminifera with complex
shell structure react in a highly specic manner to the different benthic environments and, therefore, the composition of the assemblages and the distribution patterns of
particular species reect simultaneously bottom types and
the light gradient. In the course of Earth history, larger
Foraminifera are replaced frequently. In particular, associations of Foraminifera characterizing particular shallow
water facies types are dying out and are replaced after a
certain time interval by new associations with the same
structure of shell morphology, emerging from a new evolutionary process of adaptation. These evolutionary processes make the larger Foraminifera useful as index fossils for the Permian, Jurassic, Cretaceous and Cenozoic.

7 Uses of foraminifera

Foraminifera Baculogypsina sphaerulata of Hatoma Island,

Japan. Field width = 5.22 mm.

Deep-sea species

Foraminifera are found in the deepest parts of the ocean

such as the Mariana Trench, including the Challenger
Deep, the deepest part known. At these depths, below
the carbonate compensation depth, the calcium carbonate of the tests is soluble in water due to the extreme
pressure. The Foraminifera found in the Challenger Deep
thus have no carbonate test, but instead have one of organic material.[23]
Four species found in the Challenger Deep are unknown
from any other place in the oceans, one of which is representative of an endemic genus unique to the region.
They are Resigella laevis and R. bilocularis, Nodellum aculeata, and Conicotheca nigrans (the unique genus). All
have tests that are mainly of transparent organic material
which have small (~ 100 nm) plates that appears to be
clay [23]

Evolutionary signicance

Dying planktonic Foraminifera continuously rain down

on the sea oor in vast numbers, their mineralized tests
preserved as fossils in the accumulating sediment. Beginning in the 1960s, and largely under the auspices
of the Deep Sea Drilling, Ocean Drilling, and International Ocean Drilling Programmes, as well as for the
purposes of oil exploration, advanced deep-sea drilling
techniques have been bringing up sediment cores bearing
Foraminifera fossils.[24] The eectively unlimited sup-

Because of their diversity, abundance, and complex morphology, fossil foraminiferal assemblages are useful for
biostratigraphy, and can accurately give relative dates to
sedimentary rocks. The oil industry relies heavily on
microfossils such as forams to nd potential hydrocarbon
deposits.[26]
Calcareous fossil Foraminifera are formed from elements
found in the ancient seas they lived in. Thus they are very
useful in paleoclimatology and paleoceanography. They
can be used to reconstruct past climate by examining
the stable isotope ratios and trace element content of the
shells (tests). Global temperature and ice volume can be
revealed by the isotopes of oxygen, and the history of the
carbon cycle and oceanic productivity by examining the
stable isotope ratios of carbon;[27] see 18O and 13C.
The concentration of trace elements, like magnesium
(Mg),[28] lithium (Li)[29] and boron (B),[30] also hold a
wealth of information about global temperature cycles,
continental weathering and the role of the ocean in the
global carbon cycle. Geographic patterns seen in the fossil records of planktonic forams are also used to reconstruct ancient ocean currents. Because certain types of
Foraminifera are found only in certain environments, they
can be used to gure out the kind of environment under
which ancient marine sediments were deposited.
For the same reasons they make useful biostratigraphic
markers, living foraminiferal assemblages have been used
as bioindicators in coastal environments, including indicators of coral reef health. Because calcium car-

bonate is susceptible to dissolution in acidic conditions,

Foraminifera may be particularly aected by changing
climate and ocean acidication.
Foraminifera have many uses in petroleum exploration
and are used routinely to interpret the ages and paleoenvironments of sedimentary strata in oil wells.[31] Agglutinated fossil Foraminifera buried deeply in sedimentary
basins can be used to estimate thermal maturity, which is
a key factor for petroleum generation. The Foraminiferal
Colouration Index [32] (FCI) is used to quantify colour
changes and estimate burial temperature. FCI data is particularly useful in the early stages of petroleum generation
(~100 C).
Foraminifera can also be utilised in archaeology in the
provenancing of some stone raw material types. Some
stone types, such as limestone, are commonly found to
contain fossilised Foraminifera. The types and concentrations of these fossils within a sample of stone can be
used to match that sample to a source known to contain
the same fossil signature.

Gallery
Foraminifera of Pag Island, Adriatic Sea 60 m,
eld width = 5.5 mm
Foraminifera of Pag Island, Adriatic Sea 60 m,
eld width = 5.5 mm
Foraminifera of Pag Island, Adriatic Sea 60 m,
eld width = 5.5 mm
Foraminifera of Pag Island, Adriatic Sea 60 m,
eld width = 5.5 mm
Foraminifera of the Indian Ocean, south-eastern
coast of Bali, eld width = 5.5 mm
Foraminifera of the Indian Ocean, south-eastern
coast of Bali, eld width = 5.5 mm
Foraminifera of the Indian Ocean, south-eastern
coast of Bali, eld width = 5.5 mm
Foraminifera in Ngapali, Myanmar, eld width =
5.22 mm
Foraminifera Heterostegina depressa, eld width =
4.4 mm

References

[1] Giere, Olav (2009). Meiobenthology: the microscopic

motile fauna of aquatic sediments (2nd ed.). Berlin:
Springer. ISBN 978-3540686576.

REFERENCES

[2] Lejzerowicz, Franck; Pawlowski, Jan; Fraissinet-Tachet,

Laurence; Marmeisse, Roland (1 September 2010).
Molecular evidence for widespread occurrence of
Foraminifera in soils. Environmental Microbiology 12
(9): 251826. doi:10.1111/j.1462-2920.2010.02225.x.
PMID 20406290.
[3] Kennett, J.P.; Srinivasan, M.S. (1983). Neogene planktonic foraminifera: a phylogenetic atlas. Hutchinson Ross.
ISBN 978-0-87933-070-5.
[4] World Foraminifera Database.
[5] Marshall M (3 February 2010). Zoologger: 'Living
beach ball' is giant single cell. New Scientist.
[6]
[7] Sen Gupta, Barun K. (2002). Modern Foraminifera.
Springer. p. 16. ISBN 978-1-4020-0598-5.
[8] Cavalier-Smith, T (2004). Only Six Kingdoms of Life
(PDF).
[9] Cavalier-Smith, T (2003). Protist phylogeny and the
high-level classication of Protozoa. European Journal
of Protistology 34 (4): 338348. doi:10.1078/0932-473900002.
[10] Tolweb Cercozoa
[11] European Register of Marine Species
[12] eForams-taxonomy
[13] Sen Gupta, Barun K. (1982). Ecology of benthic
Foraminifera. In Broadhead, T.W. Foraminifera: notes
for a short course organized by M.A. Buzas and B.K.
Sen Gupta. Studies in Geology 6. University of Tennessee, Dept. of Geological Sciences. pp. 3750. ISBN
0910249059. OCLC 9276403.
[14] Hemleben, C.; Anderson, O.R.; Spindler, M. (1989).
Modern Planktonic Foraminifera. Springer-Verlag. ISBN
978-3-540-96815-3.
[15] Kucera, M.; Darling, K.F. (April 2002). Cryptic species
of planktonic foraminifera: their eect on palaeoceanographic reconstructions. Philos Trans A Math Phys Eng
Sci 360 (1793): 695718. doi:10.1098/rsta.2001.0962.
PMID 12804300.
[16] Bernhard, J. M.; Bowser, S.M. (1999). Benthic
Foraminifera of dysoxic sediments: chloroplast sequestration and functional morphology. Earth Science Reviews 46: 149165. Bibcode:1999ESRv...46..149B.
doi:10.1016/S0012-8252(99)00017-3.
[17] Moore, R.C.; Lalicker, A.G.; Fischer, C.G. (1952). Ch
2 Foraminifera and Radiolaria. Invertebrate Fossils.
McGraw-Hill. OCLC 547380.
[18] Lana, C (2001). Cretaceous Carterina (Foraminifera)".
Marine Micropaleontology 41: 97. doi:10.1016/S03778398(00)00050-5.
[19] Dartnell L (8 May 2008). Sea creatures had a thing for
bling. New Scientist (2655).

[20] Foraminifera: History of Study, University College London, retrieved 20 September 2007

10 External links

[21] Langer, M. R.; Silk, M. T. B.; Lipps, J. H. (1997). Global

ocean carbonate and carbon dioxide production: The role
of reef Foraminifera. Journal of Foraminiferal Research
27 (4): 271277. doi:10.2113/gsjfr.27.4.271.

General information

[22] Adl, S. M.; Simpson, A. G. B.; Farmer, M. A.; Anderson et al. (2005). The new higher level classication
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. PMID
16248873.
[23] Gooday, A.J.; Todo, Y.; Uematsu, K.; Kitazato, H.
(July 2008). New organic-walled Foraminifera (Protista) from the oceans deepest point, the Challenger Deep
(western Pacic Ocean)". Zoological Journal of the Linnean Society 153 (3): 399423. doi:10.1111/j.10963642.2008.00393.x.
[24] http://www.nature.com/nature/debates/fossil/fossil_1.
html
[25] Journal bioinformatics and biology insights, Using the
Multiple Analysis Approach to Reconstruct Phylogenetic Relationships among Planktonic Foraminifera from
Highly Divergent and Length-polymorphic SSU rDNA
Sequences
[26] Boardman, R.S.; Cheetham, A.H.; Rowell, A.J. (1987).
Fossil Invertebrates. Wiley. ISBN 0865423024.
[27] Zachos, J.C.; Pagani, M.; Sloan, L.; Thomas, E.;
Billups, K. (2001). Trends, Rhythms, and Aberrations in Global Climate, 65 Ma to Present. Science
292 (5517): 686693. Bibcode:2001Sci...292..686Z.
doi:10.1126/science.1059412. PMID 11326091.
[28] Branson, Oscar; Redfern, Simon A.T.; Tyliszczak,
Tolek; Sadekov, Aleksey; Langer, Gerald; Kimoto, Katsunori; Eldereld, Henry (December
2013).
The coordination of Mg in foraminiferal
calcite.
Earth and Planetary Science Letters
383:
134141.
Bibcode:2013E&PSL.383..134B.
doi:10.1016/j.epsl.2013.09.037.
[29] Misra, S.; Froelich, P. N. (26 January 2012). Lithium
Isotope History of Cenozoic Seawater: Changes in Silicate Weathering and Reverse Weathering. Science 335
(6070): 818823. doi:10.1126/science.1214697. PMID
22282473.
[30] Hemming, N.G.; Hanson, G.N. (January 1992). Boron
isotopic composition and concentration in modern marine
carbonates. Geochimica et Cosmochimica Acta 56 (1):
537543. doi:10.1016/0016-7037(92)90151-8.
[31] Jones, R.W. (1996). Micropalaeontology in petroleum exploration. Clarendon Press. ISBN 978-0-19-854091-5.
[32] McNeil, D.H.; Issler, D.R.; Snowdon, L.R. (1996).
Colour Alteration, Thermal Maturity, and Burial Diagenesis in Fossil Foraminifers. Geological Survey of Canada
Bulletin 499. Geological Survey of Canada. ISBN 9780-660-16451-9.

The University of California Museum of Paleontology website has an Introduction to the Foraminifera
Researchers at the University of South Florida developed a system using Foraminifera for monitoring
coral reef environments
University College Londons micropaleontology site
has an overview of Foraminifera, including many
high-quality SEMs
Illustrated glossary of terms used in foraminiferal
research is the Lukas Hottingers glossary published
in the OA e-journal Carnets de Gologie Notebooks on Geology
Information on Foraminifera Martin Langers Micropaleontology Page
Benthic Foraminifera information from the 2005
Urbino Summer School of Paleoclimatology
Online ip-books
Illustrated glossary of terms used in foraminiferal
research by Lukas Hottinger (alternative version of
the one published in Carnets de Gologie Notebooks on Geology)
Resources
The star*sand project (part of micro*scope)
is a cooperative database of information about
Foraminifera
3D models of forams, generated by X-ray tomography
CHRONOS has several Foraminifera resources, including a taxon search page and a micro-paleo section
eForams is a web site focused on Foraminifera and
modeling of foraminiferal shells
Foraminifera Gallery Illustrated catalog of recent
and fossil Foraminifera by genus and locality
"Foraminifera". NCBI Taxonomy Browser. 29178.

11

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Foraminifera Source: http://en.wikipedia.org/wiki/Foraminifera?oldid=665287797 Contributors: Josh Grosse, Azhyd, Dcljr, CatherineMunro, Kils, Denis Barthel, Topbanana, Hadal, UtherSRG, Lproven, Guanaco, Mboverload, Tipiac, Espetkov, Ukexpat,
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