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Temporal range: 542–0  Ma [1]
Ammonia tepida.jpg
Live Ammonia tepida (Rotaliida)
Scientific classification

d'Orbigny, 1826

incertae sedis


Foraminifera ( /fəˌræməˈnɪfərə/ ; Latin for "hole bearers"; informally called "forams") are members of a phylum or class of amoeboid protists characterized by streaming granular ectoplasm for catching food and other uses; and commonly an external shell (called a "test") of diverse forms and materials. Tests of chitin (found in some simple genera, and Textularia in particular) are believed to be the most primitive type. Most foraminifera are marine, the majority of which live on or within the seafloor sediment (i.e., are benthic), while a smaller variety float in the water column at various depths (i.e., are planktonic). Fewer are known from freshwater or brackish conditions, and some very few (nonaquatic) soil species have been identified through molecular analysis of small subunit ribosomal DNA. [2] [3]

Latin Indo-European language of the Italic family

Latin is a classical language belonging to the Italic branch of the Indo-European languages. The Latin alphabet is derived from the Etruscan and Greek alphabets and ultimately from the Phoenician alphabet.

Amoeba polyphyletic group including different eucariot taxons

An amoeba, often called amoeboid, is a type of cell or unicellular organism which has the ability to alter its shape, primarily by extending and retracting pseudopods. Amoebas do not form a single taxonomic group; instead, they are found in every major lineage of eukaryotic organisms. Amoeboid cells occur not only among the protozoa, but also in fungi, algae, and animals.

Protist unicellular organism of a diverse group of eukaryotic microorganisms

A protist is any eukaryotic organism that is not an animal, plant or fungus. The protists do not form a natural group, or clade, since they exclude certain eukaryotes; but, like algae or invertebrates, they are often grouped together for convenience. In some systems of biological classification, such as the popular five-kingdom scheme proposed by Robert Whittaker in 1969, the protists make up a kingdom called Protista, composed of "organisms which are unicellular or unicellular-colonial and which form no tissues".

Foraminifera typically produce a test, or shell, which can have either one or multiple chambers, some becoming quite elaborate in structure. [4] These shells are commonly made of calcium carbonate (CaCO
) or agglutinated sediment particles. Over 50,000 species are recognized, both living (10,000) [5] and fossil (40,000). [6] [7] They are usually less than 1 mm in size, but some are much larger, the largest species reaching up to 20 cm. [8]

Test (biology) hard shell of some spherical marine animals, notably sea urchins and microorganisms such as testate foraminiferans, radiolarians, and testate amoebae

In biology, a test is the hard shell of some spherical marine animals, notably sea urchins and microorganisms such as testate foraminiferans, radiolarians, and testate amoebae.

Calcium carbonate Chemical compound

Calcium carbonate is a chemical compound with the formula CaCO3. It is a common substance found in rocks as the minerals calcite and aragonite (most notably as limestone, which is a type of sedimentary rock consisting mainly of calcite) and is the main component of pearls and the shells of marine organisms, snails, and eggs. Calcium carbonate is the active ingredient in agricultural lime and is created when calcium ions in hard water react with carbonate ions to create limescale. It is medicinally used as a calcium supplement or as an antacid, but excessive consumption can be hazardous.

Agglutination is the clumping of particles. The word agglutination comes from the Latin agglutinare.

In modern Scientific English, the term foraminifera is both singular and plural (irrespective of the word's Latin derivation), and is used to describe one or more specimens or taxa: its usage as singular or plural must be determined from context. Foraminifera is frequently used informally to describe the group, and in these cases is generally lowercase. [9]


The taxonomic position of the Foraminifera has varied since their recognition as protozoa (protists) by Schultze in 1854, [10] there referred to as an order, Foraminiferida. Loeblich and Tappan (1992) reranked Foraminifera as a class [11] as it is now commonly regarded.

Alfred R. ("Al") Loeblich Jr (1914–1994) was an American micropaleontologist. He was married to Helen Niña Tappan Loeblich and the two co-authored a number of important works on the Foraminifera and related organisms.

Helen Niña Tappan Loeblich was a leading micropaleontologist, a professor of geology at the University of California, Los Angeles, a United States Geological Survey (USGS) biostratigrapher, and a scientific illustrator whose micropaleontology specialty was research on Cretaceous foraminifera.

The Foraminifera have typically been included in the Protozoa, [12] [13] [14] or in the similar Protoctista or Protist kingdom. [15] [16] Compelling evidence, based primarily on molecular phylogenetics, exists for their belonging to a major group within the Protozoa known as the Rhizaria. [12] Prior to the recognition of evolutionary relationships among the members of the Rhizaria, the Foraminifera were generally grouped with other amoeboids as phylum Rhizopodea (or Sarcodina) in the class Granuloreticulosa.

Protozoa Diverse motile unicellular heterotrophic eukaryotic organisms

Protozoa is an informal term for single-celled eukaryotes, either free-living or parasitic, which feed on organic matter such as other microorganisms or organic tissues and debris. Historically, the protozoa were regarded as "one-celled animals", because they often possess animal-like behaviors, such as motility and predation, and lack a cell wall, as found in plants and many algae. Although the traditional practice of grouping protozoa with animals is no longer considered valid, the term continues to be used in a loose way to identify single-celled organisms that can move independently and feed by heterotrophy.

In biology, kingdom is the second highest taxonomic rank, just below domain. Kingdoms are divided into smaller groups called phyla.

Molecular phylogenetics The branch of phylogeny that analyzes genetic, hereditary molecular differences

Molecular phylogenetics is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominately in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to determine the processes by which diversity among species has been achieved. The result of a molecular phylogenetic analysis is expressed in a phylogenetic tree. Molecular phylogenetics is one aspect of molecular systematics, a broader term that also includes the use of molecular data in taxonomy and biogeography.

The Rhizaria are problematic, as they are often called a "supergroup", rather than using an established taxonomic rank such as phylum. Cavalier-Smith defines the Rhizaria as an infra-kingdom within the kingdom Protozoa. [12]

Taxonomic rank Level in a taxonomic hierarchy

In biological classification, taxonomic rank is the relative level of a group of organisms in a taxonomic hierarchy. Examples of taxonomic ranks are species, genus, family, order, class, phylum, kingdom, domain, etc.

In biology, a phylum is a level of classification or taxonomic rank below kingdom and above class. Traditionally, in botany the term division has been used instead of phylum, although the International Code of Nomenclature for algae, fungi, and plants accepts the terms as equivalent. Depending on definitions, the animal kingdom Animalia or Metazoa contains approximately 35 phyla, the plant kingdom Plantae contains about 14, and the fungus kingdom Fungi contains about 8 phyla. Current research in phylogenetics is uncovering the relationships between phyla, which are contained in larger clades, like Ecdysozoa and Embryophyta.

Some taxonomies put the Foraminifera in a phylum of their own, putting them on par with the amoeboid Sarcodina in which they had been placed.

Although as yet unsupported by morphological correlates, molecular data strongly suggest the 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. [13] However, the exact relationships of the forams to the other groups and to one another are still not entirely clear. Foraminifera are closely related to testate amoebae. [17]

The most recent taxonomy by Mikhalevich 2013. [18]

Living Foraminifera

Modern Foraminifera are primarily marine organisms, but living individuals have been found in brackish, freshwater [19] and even terrestrial habitats. [3] The majority of the species are benthic, and a further 40 morphospecies are planktonic. [20] This count may, however, represent only a fraction of actual diversity, since many genetically distinct species may be morphologically indistinguishable. [21]

A number of forams have unicellular algae as endosymbionts, from diverse lineages such as the green algae, red algae, golden algae, diatoms, and dinoflagellates. [20] Some forams are kleptoplastic, retaining chloroplasts from ingested algae to conduct photosynthesis. [22]


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. [19] The pseudopods are used for locomotion, anchoring, and in capturing food, which consists of small organisms such as diatoms or bacteria. [20]

The generalized foraminiferal life-cycle involves an alternation between haploid and diploid generations, although they are mostly similar in form. [10] [23] 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 divides to produce new gamonts. Multiple rounds of asexual reproduction between sexual generations are not uncommon in benthic forms. [19]


Foraminiferan tests (ventral view) Benthic foraminifera.jpg
Foraminiferan tests (ventral view)
Fossil nummulitid foraminiferans showing microspheric and megalospheric individuals; Eocene of the United Arab Emirates; scale in mm Nummulitids.jpg
Fossil nummulitid foraminiferans showing microspheric and megalospheric individuals; Eocene of the United Arab Emirates; scale in mm
The miliolid foraminiferan Quinqueloculina from the North Sea Quinqueloculina seminula.jpg
The miliolid foraminiferan Quinqueloculina from the North Sea
Thin section of a peneroplid foraminiferan from Holocene lagoonal sediment in Rice Bay, San Salvador Island, Bahamas. Scale bar 100 micrometres Peneroplid thin section PP.jpg
Thin section of a peneroplid foraminiferan from Holocene lagoonal sediment in Rice Bay, San Salvador Island, Bahamas. Scale bar 100 micrometres
Ammonia beccarii, a benthic foram from the North Sea. Ammonia beccarii.jpg
Ammonia beccarii , a benthic foram from the North Sea.
Foraminifera Baculogypsina sphaerulata of Hatoma Island, Japan. Field width 5.22 mm 2085f Japon Hatoma.jpg
Foraminifera Baculogypsina sphaerulata of Hatoma Island, Japan. Field width 5.22 mm

The form and composition of their tests are the primary means by which forams are identified and classified. Most have calcareous tests, composed of calcium carbonate. [19] In other forams, the tests 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. The test contains an organic matrix, which can sometimes be recovered from fossil samples. [24]

Tests as fossils are known from as far back as the Cambrian period, [25] 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. [26] It is estimated that reef Foraminifera generate about 43 million tons of calcium carbonate per year. [27]

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. [28]

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. [29]

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 (about 100 nm) plates that appear to be clay. [29]

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. [30] 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. [30] 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 [31] 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. [32] These evolutionary processes make the larger Foraminifera useful as index fossils for the Permian, Jurassic, Cretaceous and Cenozoic.


Because of their diversity, abundance, and complex morphology, fossil foraminiferal assemblages are useful for biostratigraphy, and can accurately give relative dates to sedimentary rocks, as was discovered by Alva C. Ellisor in 1920. [33] The oil industry relies heavily on microfossils such as forams to find potential hydrocarbon deposits. [34]

Calcareous fossil Foraminifera are formed from elements found in the ancient seas where they lived. Thus, they are very useful in paleoclimatology and paleoceanography. They can be used, as a climate proxy, 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; [35] see δ18O and δ13C. The concentration of trace elements, like magnesium (Mg), [36] lithium (Li) [37] and boron (B), [38] 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 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 have many uses in petroleum exploration and are used routinely to interpret the ages and paleoenvironments of sedimentary strata in oil wells. [39] 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 [40] (FCI) is used to quantify colour changes and estimate burial temperature. FCI data is particularly useful in the early stages of petroleum generation (about 100 °C).

Foraminifera can also be used 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".

Related Research Articles

Radiolaria phylum of protists

The Radiolaria, also called Radiozoa, are protozoa of diameter 0.1–0.2 mm that produce intricate mineral skeletons, typically with a central capsule dividing the cell into the inner and outer portions of endoplasm and ectoplasm.The elaborate mineral skeleton is usually made of silica. They are found as zooplankton throughout the ocean, and their skeletal remains make up a large part of the cover of the ocean floor as siliceous ooze. Due to their rapid turn-over of species, they represent an important diagnostic fossil found from the Cambrian onwards. Some common radiolarian fossils include Actinomma, Heliosphaera and Hexadoridium.

Paleocene–Eocene Thermal Maximum rapid (in geological terms) global warming, profound changes in ecosystems, and major perturbations in the carbon cycle which started about 55.0 million years ago

The Paleocene–Eocene Thermal Maximum (PETM), alternatively "Eocene thermal maximum 1" (ETM1), and formerly known as the "Initial Eocene" or "Late Paleocene Thermal Maximum", was a time period with more than 8 °C warmer global average temperature than today. This climate event began at the time boundary of the Paleogene, between the Paleocene and Eocene geological epochs. The exact age and duration of the event is uncertain but it is estimated to have occurred around 55.5 million years ago.

Xenophyophore class of amoeboid protists

Xenophyophores are multinucleate unicellular organisms found on the ocean floor throughout the world's oceans, at depths of 500 to 10,600 metres. They are a kind of foraminiferan that extract minerals from their surroundings and use them to form an exoskeleton known as a test.

Rhizaria infrakingdom of protists

The Rhizaria are a species-rich supergroup of mostly unicellular eukaryotes. A multicellular form has also been described. This supergroup was proposed by Cavalier-Smith in 2002. Being described mainly from rDNA sequences, they vary considerably in form, having no clear morphological distinctive characters (synapomorphies), but for the most part they are amoeboids with filose, reticulose, or microtubule-supported pseudopods. Many produce shells or skeletons, which may be quite complex in structure, and these make up the vast majority of protozoan fossils. Nearly all have mitochondria with tubular cristae.

The Allogromiida is an order of single-chambered, mostly organic-walled foraminiferans, including some that produce agglutinated tests (Lagynacea). Genetic studies indicate that some foraminiferans with agglutinated tests, previously included in the Textulariida or as their own order Astrorhizida, may also belong here. Allogromiids produce relatively simple tests, usually with a single chamber, similar to those of other protists such as Gromia. They are found as both marine and freshwater forms, and are the oldest forams known from the fossil record.

Fusulinida order of foraminifera (fossil)

The Fusulinida is an extinct order within the Foraminifera in which the tests (shells) are composed of tightly packed, secreted microgranular calcite. Like all Forams, they are single-celled organisms. In advanced forms the test wall is differentiated into two or more layers. Loeblich and Tappan, 1988, gives a range from the Lower Silurian to the Upper Permian, with the fusulinid foraminifera going extinct with the Permian–Triassic extinction event. While the latter is true, a more supported projected timespan is from the Mid-Carboniferous period.

Rotaliida order of foraminifers

The Rotaliida are an order of Foraminifera, characterized by multilocular tests (shells) composed of bilammelar perforate hyaline lamellar calcite that may be optically radial or granular.

The Komokiacea are a small group of amoeboid protozoa, considered to be foraminifera, though there have been suggestions that they are a separate group, closely related to foraminifera. Komokiacea are rather large organisms, often exceeding 300 micrometers in maximum dimensions. Along with Xenophyophores they dominate the macro- and megabenthic fauna in the deep sea and are commonly referred to as "giants protists".

Miliolacea superfamily of protists

Miliolacea is one of five superfamilies belonging to the Miliolida,.

<i>Globigerina bulloides</i> species of foraminifer

Globigerina bulloides is a species of heterotrophic planktonic foraminifer with a wide distribution in the photic zone of the world's oceans. It is able to tolerate a range of sea surface temperatures, salinities and water densities, and is most abundant at high southern latitudes, certain high northern latitudes, and in low-latitude upwelling regions. The density or presence of G. bulloides may change as a function of phytoplankton bloom successions, and they are known to be most abundant during winter and spring months.

Globigerinoidea superfamily of foraminifers

Globigerinoidea is a superfamily of free-living, calcareous, planktonic foraminiferal protists that have lived in the open ocean since the Eocene. It is part of the suborder Globigerinina.

The Silicoloculinida are an order of Cenozoic benthic foraminifera with tests made of imperforate opaline silica. The order is known from a single genus, Miliammellus, in the family Silicoloculinidae.

<i>Cibicides</i> genus of foraminifers

Cibicides is a genus of cosmopolitan benthic foraminifera known from at least as far back as the Paleocene that extends down to the present.

Rotalidia comprises a class of Foraminifera where Foraminifera is regarded as a phylum, that unites Foraminifera that have tests composed of secreted lamellar calcium carbonate, optically radial or granular calcite, or aragonite; separating them from those with porcelaneous, agglutinated, or microgranular, tests, or tests composed of organic compounds. Seven orders are included, the:

Kingdoms animal, plant and fungi are in bold. Protists are a large and diverse group of eukaryotic microorganisms which belong to the kingdom Protista.

<i>Amphistegina</i> genus of foraminifers

Amphistegina is a genus of foraminiferal protists included in the Rotaliida with a stritigraphic range extending from the Eocene to recent and a cosmopolitan distribution. The test is an asymmetrically biconvex trochospiral that may be bi-involute or partially evolute on the spiral side. Chambers are numerous, broad. and low, strongly curved back at the periphery to form chamber prolongations. The umbilical side is stellate, like that of Asterigerina, and has a distinct umbilical plug. The wall is calcareous, optically radial; the surfacce finely perforate and smooth overall. The periphery angular to carinate (keeled); the aperture an interiomarginal slit on the umbilical side, bordered by a lip.

Miliollata is a class wherein Foraminifera is regarded as a phylum that unites the porcelaneous Miliolida, and siliceous Rzehakinidae based on similarities of their tests. Previously the Rzehakinidae were included in the Textulariina based on test wall composition rather than test form. The meaning of Miliolida is retained.

Rotaliana is a subclass of benthic Foraminifera with multichambered tests of perforate hyaline calcite. Tests may be planospiral, low or high trochospiral, or serial. Interiors may be complex with secondary chambers and interconnecting canal systems. Rotaliana are separate from the planktonic Globigerinana although both have tests of similar composition. The Textulariana, which contains forms that are rather similar, differs in be agglutinated.

Globigerinana are free living pelagic foraminiferan, included in the class Rotaliata that range from the Jurassic to recent. Test are commonly planospiral or trochospiral but may be uniserial to multiserial and are of secreted hyaline (glassy) calcite. Chambers are flattned in planospiral forms and spheroidal in trochospiral and serial forms. Some have long radial spines, or needles that may be solidly fixed or moveable in sockets. Gametes are biflagelate and are produced are greater number than by bottom dwelling benthic forms.

Chilostomelloidea is a superfamily of foraminifera in the order Rotaliida. They are found in sediments of Early Cretaceous (Barremian) to the present.


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