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Foraminifera
Fossil range: Precambrian - Recent
Live Ammonia tepida (Rotaliida)
Scientific classification
Domain: Eukaryota
Kingdom: Rhizaria
Superphylum: Retaria
Phylum: Foraminifera
d'Orbigny, 1826
Orders

Allogromiida
Carterinida
Fusulinida - extinct
Globigerinida
Involutinida - extinct
Lagenida
Miliolida
Silicoloculinida
Spirillinida
Textulariida
incertae sedis
   Xenophyophorea
   Reticulomyxa

The Foraminifera, ("Hole Bearers") or forams for short, are a large group of amoeboid protists with reticulating pseudopods, fine strands of cytoplasm that branch and merge to form a dynamic net.[1] They typically produce a test, or shell, which can have either one or multiple chambers, some becoming quite elaborate in structure.[2] These shells are made of calcium carbonate (CaCO3) or agglutinated sediment particles. About 275,000 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.

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.[3] However, the exact relationships of the forams to the other groups and to one another are still not entirely clear.

Contents

Living forams

Modern forams are primarily marine, although some can survive in brackish conditions.[4] A few species survive in fresh water and one even lives in damp rainforest soil. They are most commonly benthic, and about 40 morphospecies are planktonic.[1] This count may however represent only a fraction of actual diversity, since many genetically discrepant species may be morphologically indistinguishable.[5]

A number of forms have unicellular algae as endosymbionts, from diverse lineages such as the green algae, red algae, golden algae, diatoms, and dinoflagellates.[1] Some forams are kleptoplastic, retaining chloroplasts from ingested algae to conduct photosynthesis.[6]

Biology

The foraminiferal cell is divided into granular endoplasm and transparent ectoplasm from which a pseudopodial net may emerge through a single opening or through many perforations in the test. Individual pseudopods characteristically have small granules streaming in both directions.[4] The pseudopods are used for locomotion, anchoring, and in capturing food, which consists of small organisms such as diatoms or bacteria.[1]

The foraminiferal life-cycle involves an alternation between haploid and diploid generations, although they are mostly similar in form.[7][8] 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 in benthic forms.[4]

Tests

Foraminiferan tests (ventral view)
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 identified and classified. Most have calcareous tests, composed of calcium carbonate.[4] 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,[9] and many marine sediments are composed primarily of them. For instance, the limestone that makes up the pyramids of Egypt is composed almost entirely of nummulitic benthic foraminifera.[10] Production estimates indicate that reef foraminifera annually generate approximately 43 million tons of calcium carbonate and thus play an essential role in the production of reef carbonates.[11]

Genetic studies have identified the naked amoeba "Reticulomyxa" and the peculiar xenophyophores as foraminiferans without tests. A few other amoeboids 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.[12]

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.[13]

Four species have been found in the Challenger Deep that are unknown from any other place in the ocean, 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 [13]

Evolutionary significance

Dying planktonic foraminifera continuously rain down on the sea floor 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 by the millions. The effectively unlimited supply 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. 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. Larger benthic foraminifera with complex shell structure react in a highly specific manner to the different benthic environments and, therefore, the composition of the assemblages and the distribution patterns of particular species reflect 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 prone to be fossil index for the Permian, Jurassic, Cretaceous and Cenozoic (e.g. Lukas Hottinger).

Uses of forams

Because of their diversity, abundance, and complex morphology, fossil foraminiferal assemblages are useful for biostratigraphy, and can accurately give relative dates to rocks. The oil industry relies heavily on microfossils such as forams to find potential oil deposits.[14]

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 of oxygen, and the history of the carbon cycle and oceanic productivity by examining the stable isotope ratios of carbon;[15] see δ18O and δ13C. 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 figure 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 carbonate is susceptible to dissolution in acidic conditions, foraminifera may be particularly affected by changing climate and ocean acidification.

Foraminifera can also be utilised in archaeology in the provenancing of some stone raw material types. Some stone types, such as chert, 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".

References

  1. ^ a b c d Hemleben, C.; Spindler, M.& Anderson, O.R. (1989). Modern Planktonic Foraminifera. Springer-Verlag. pp. 363. 
  2. ^ Kennett, J.P.; Srinivasan, M.S. (1983). Neogene Planktonic Foraminifera: A Phylogenetic Atlas. Hutchinson Ross. pp. 265. 
  3. ^ Cavalier-Smith, T. (2003). "Protist phylogeny and the high-level classification of Protozoa". European Journal of Protistology 34 (4): 338–348. doi:10.1078/0932-4739-00002. 
  4. ^ a b c d Sen Gupta, B.K. (1983). Modern Foraminifera. Springer. pp. 384. 
  5. ^ Kucera, M.; Darling, K.F. (2002). "Genetic diversity among modern planktonic foraminifer species: its effect on paleoceanographic reconstructions". Philosophical Transactions of the Royal Society of London A360 (4): 695–718. 
  6. ^ Bernhard, J. M.; Bowser, S.M. (1999). "Benthic foraminifera of dysoxic sediments: chloroplast sequestration and functional morphology". Earth Science Reviews 46: 149–165. doi:10.1016/S0012-8252(99)00017-3. 
  7. ^ Moore, Lalicker, and Fischer. Invertebrate Fossils, Ch 2 Foraminifera and Radiolaria. McGraw-Hill 1952
  8. ^ Loeblich, A and Tappan H. Foraminiferida in the Treatise on Invertebrate Paleontology Part C Protista 2 Sarcodina; Geological Society of America and University of Kansas Press, 1964, fifth printing 1984
  9. ^ Sea creatures had a thing for bling - life - 08 May 2008 - New Scientist
  10. ^ Foraminifera: History of Study, University College London, retrieved 20 September 2007
  11. ^ 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): 271–277. doi:10.2113/gsjfr.27.4.271. 
  12. ^ Adl, SM; Simpson, AG; Farmer, MA; Andersen, RA; Anderson, OR; Barta, JR; Bowser, SS; Brugerolle, G et al. (2005). "The new higher level classification 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. 
  13. ^ a b New organic-walled Foraminifera (Protista) from the ocean's deepest point, the Challenger Deep (western Pacific Ocean) A. J. Gooday, Y. Todo, K. Uematsu, and H. Kitazato, Zoological Journal of the Linnean Society, 2008, 153, 399–423.
  14. ^ Boardman, R.S. (1987). Fossil Invertebrates. Blackwell. pp. 714. 
  15. ^ Zachos, J.C.; Pagani, M., Sloan, L., Thomas, E., and Billups, K. (2001). "Trends, Rhythms, and Aberrations in Global Climate, 65 Ma to Present". Science 292 (5517): 686–693. doi:10.1126/science.1059412. PMID 11326091. 

External links

General information
Online flip-Books

Illustrated glossary of terms used in foraminiferal research by Lukas Hottinger (alternative version of the one published in "Carnets de Géologie - Notebooks on Geology")

Resources
  • eForams is a web site focused on foraminifera and modeling of foraminiferal shells


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