Foraminifera

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{{Taxobox | color = khaki | name = Foraminifera | fossil_range = Cambrian - Recent | image = live_Ammonia_tepida.jpg | image_width = 200px | image_caption = Live Ammonia tepida (Rotaliida) | regnum = Protista | phylum = Foraminifera | subdivision_ranks = Orders | subdivision = Allogromiida
Carterinida
Fusulinida - extinct
Globigerinida
Involutinida - extinct
Lagenida
Miliolida
Robertinida
Rotaliida
Silicoloculinida
Spirillinida
Textulariida
incertae sedis
   Xenophyophorea
   Reticulomyxa }} The Foraminifera, or forams for short, are a large group of amoeboid protists with reticulating pseudopods, fine strands that branch and merge to form a dynamic net. They typically produce a shell, or test, which can have either one or multiple chambers, some becoming quite elaborate in structure. About 250 000 species are recognized, both living and fossil. They are usually less than 1 mm in size, but some are much larger, and the largest recorded specimen reached 19 cm. As fossils, foraminifera are extremely useful

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Living forams

Modern forams are primarily marine, although they can survive in brackish conditions. A few species survive in fresh water (e.g. Lake Geneva) and one species even lives in damp rainforrest soil. They are very common in the meiobenthos, and about 40 species are planktonic. The cell is divided into granular endoplasm and transparent ectoplasm. The pseudopodial net may emerge through a single opening or many perforations in the test, and characteristically has small granules streaming in both directions.

The pseudopods are used for locomotion, anchoring, and in capturing food, which consists of small organisms such as diatoms or bacteria. A number of forms have unicellular algae as endosymbionts, from diverse lineages such as the green algae, red algae, golden algae, diatoms, and dinoflagellates. Some forams are kleptoplastic, retaining chloroplasts from ingested algae to conduct photosynthesis.

The foraminiferan life-cycle involves an alternation between haploid and diploid generations, although they are mostly similar in form. The haploid or gamont initially has a single nucleus, and divides to produce numerous gametes, which typically have two flagella. The diploid or schizont is multinucleate, and after meiosis fragments to produce new gamonts. Multiple rounds of asexual reproduction between sexual generations is not uncommon.

Image:Benthic foraminifera.jpg

Tests

The form and composition of the test is the primary means by which forams are identified and classified. Most have calcareous tests, composed of calcium carbonate, which generally takes the form of interlocking microscopic crystals, giving it a glassy or hyaline appearance. 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 flow between chambers, are called apertures.

Tests are known as fossils as far back as the Cambrian period, and many marine sediments are composed primarily of them. For instance, the nummulitic limestone that makes up the pyramids of Egypt is composed almost entirely of them. Forams may also make a significant contribution to the overall deposition of calcium carbonate in coral reefs.

Utility of foraminifera

Because of their diversity, abundance, and complex morphology, fossil foraminiferal assembleages can give accurate relative dates for rocks and thus are extremely useful in biostratigraphy. Before more modern techniques became available, the oil industry relied heavily on microfossils such as foraminifera to find potential oil deposits.

For the same reasons they make good biostratigraphic markers, living foraminiferal assembleages have been used as bioindicators in coastal environments, including as indicators of coral reef health.

Fossil foraminifera are also useful in paleoclimatology and paleoceanography. They can be used to reconstruct past climate by examining their oxygen stable isotope ratios. Geographic patterns seen in the fossil record of planktonic forams are also used to reconstruct paleo ocean current patterns.


Relationships

Genetic studies have identified the naked amoeba Reticulomyxa and the peculiar xenophyophores as foraminiferans without tests. A few other ameoboids produce reticulose pseudopods, and were formerly classified with the forams as the Granuloreticulosa, but this is no longer considered a natural group, and most are now placed among the Cercozoa. Both the Cercozoa and Radiolaria are close relatives of the Foraminifera, together making up the Rhizaria, but the exact position of the forams is still unclear.

See also

External links

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