Foraminifera

Foraminifera (/fəˌræməˈnɪfərə/; Latin for "hole bearers";

informally called "forams") are single-celled organisms, members Foraminifera
of a phylum or class of amoeboid protists characterized by Temporal range: 542–0 Ma [1]
streaming granular ectoplasm for catching food and other uses; and
PreꞒ Ꞓ O S D C P T J K PgN
commonly an external shell (called a "test") of diverse forms and
Ediacaran–Recent
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
number 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]
Live Ammonia tepida (Rotaliida)
Foraminifera typically produce a test, or shell, which can have
either one or multiple chambers, some becoming quite elaborate in Scientific classification
structure.[4] These shells are commonly made of calcium carbonate Domain: Eukaryota
(CaCO3 ) or agglutinated sediment particles. Over 50,000 species
(unranked): SAR
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, (unranked): Rhizaria
the largest species reaching up to 20 cm.[8]
Phylum: Retaria
In modern scientific English, the term foraminifera is both singular Subphylum: Foraminifera
and plural (irrespective of the word's Latin derivation), and is used
d'Orbigny, 1826
to describe one or more specimens or taxa: its usage as singular or
plural must be determined from context. Foraminifera is frequently Subdivisions
used informally to describe the group, and in these cases is
generally lowercase.[9] "Monothalamea"

"Allogromiida"
Contents "Astrorhizida"
History of study Xenophyophorea
Taxonomy Reticulomyxa
Biology
Tubothalamea
Reproduction
Variations in reproductive mode Miliolida
Tests Spirillinida
Test composition
Soft tests Silicoloculinida
Agglutinated tests
Globothalamea
Calcareous tests
Silica tests
 Test wall construction Textulariida
Deep-sea species Rotaliida
Evolutionary history
Globigerinida
Paleontological applications
Carterinida
Modern uses
Robertinida
Gallery
References Fusulinida? — extinct

External links incertae sedis

Involutinida
History of study
Lagenida
The earliest known reference to foraminifera comes from
Herodotus, who in the 5th century BCE noted them as making up the rock that forms the Great Pyramid of
Giza. These are today recognized as representatives of the genus Nummulites. Strabo, in the 1st Century BCE,
noted the same foraminifera, and suggested that they were the remains of lentils left by the workers who built
the pyramids.[10]

Robert Hooke observed a foraminifera under the microscope, as described and illustrated in his 1665 book
Micrographia:

I was trying several small and single Magnifying Glasses, and casually viewing a parcel of white
Sand, when I perceiv'd one of the grains exactly shap'd and wreath'd like a Shell[...] I view'd it
every way with a better Microscope and found it on both sides, and edge-ways, to resemble the
Shell of a small Water-Snail with a flat spiral Shell[...][11]

Antonie van Leeuwenhoek described and illustrated foraminiferal tests in 1700, describing them as minute
cockles; his illustration is recognizable as being Elphidium.[12] Early workers classified foraminifera within the
genus Nautilus, noting their similarity to certain cephalopods. It was recognised by Lorenz Spengler in 1781
that foraminifera had holes in the septa, which would eventually grant the group its name.[13] Spengler also
noted that the septa of foraminifera arced the opposite way from those of nautili and that they lacked a nerve
tube.[14]

Alcide d'Orbigny, in his 1826 work, considered them to be a group of minute cephalopods and noted their odd
morphology, interpreting the pseudopodia as tentacles and noting the highly reduced (in actuality, absent)
head.[15] He named the group foraminifères, or "hole-bearers", as members of the group had holes in the
divisions between compartments in their shells, in contrast to nautili or ammonites.[9]

The protozoan nature of foraminifera was first recognized by Dujardin in 1835.[13] Shortly after, in 1852,
d'Orbigny produced a classification scheme, recognising 72 genera of foraminifera, which he classified based
on test shape—a scheme that drew severe criticism from colleagues.[12]

H.B. Brady's 1884 monograph described the foraminiferal finds of the Challenger expedition. Brady
recognized 10 families with 29 subfamilies, with little regard to stratigraphic range; his taxonomy emphasized
the idea that multiple different characters must separate taxonomic groups, and as such placed agglutinated and
calcareous genera in close relation.
This overall scheme of classification would remain until Cushman's
work in the late 1920s. Cushman viewed wall composition as the
single most important trait in classification of foraminifera; his
classification became widely accepted but also drew criticism from
colleagues for being "not biologically sound".

Cushman's scheme nevertheless remained the dominant scheme of

classification until Tappan and Loeblich's 1964 classification, which
placed foraminifera into the general groupings still used today, based
on microstructure of the test wall.[12] These groups have been
variously moved around according to different schemes of higher-
level classification. Pawlowski's (2013) use of molecular systematics
has generally confirmed Tappan and Loeblich's groupings, with some
being found as polyphyletic or paraphyletic; this work has also helped
to identify higher-level relationships among major foraminiferal Earliest known illustration of a
groups.[16] foraminifera shell, published by
Robert Hooke in his 1665 book
Taxonomy Micrographia.

The taxonomic position of the Foraminifera has varied since Schultze

in 1854,[17] who referred to as an order, Foraminiferida. Loeblich and Tappan (1992) reranked Foraminifera as
a class[18] as it is now commonly regarded.

The Foraminifera have typically been included in the Protozoa,[19][20][21] or in the similar Protoctista or Protist
kingdom.[22][23] Compelling evidence, based primarily on molecular phylogenetics, exists for their belonging
to a major group within the Protozoa known as the Rhizaria.[19] 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.

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.[19]

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.[20] 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.[24]
 Taxonomy from Mikhalevich 2013[25]
* Foraminifera d'Orbigny 1826

Order Reticulomyxida
Class Schizocladea Cedhagen & Mattson 1992

Order Schizocladida
Class Xenophyophorea Schultze 1904

Order Stannomida Tendal 1972

Order Psamminida Tendal 1972
Class Astrorhizata Saidova 1981

Subclass Lagynana Mikhalevich 1980

Order Ammoscalariida Mikhalevich 1980

Order Lagynida Mikhalevich 1980
Order Allogromiida Loeblich & Tappan 1961
Subclass Astrorhizana Saidova 1981

Order Astrorhizida Lankester 1885

Order Dendrophryida Mikhalevich 1995
Order Hippocrepinida Saidova 1981
Order †Parathuramminida Mikhalevich 1980
Order Psammosphaerida Haeckel 1894
Class Rotaliata Mikhalevich 1980 (hyaline foraminifers)

Subclass Globigerinana Mikhalevich 1980

Order Cassigerinellida Mikhalevich 2013

Order Globigerinida Carpenter, Parker & Jones 1862
Order Hantkeninida Mikhalevich 1980
Order Heterohelicida Fursenko 1958
Order Globorotaliida Mikhalevich 1980
Subclass Textulariana Mikhalevich 1980

Order Nautiloculinida Mikhalevich 2003

Order Spiroplectamminida Mikhalevich 1992
Order Textulariida Delage & Hérouard 1896
Order Trochamminida Saidova 1981 (Carterinida Loeblich & Tappan 1955]
Order Verneuilinida Mikhalevich & Kaminski 2003
Subclass Rotaliana Mikhalevich 1980

Superorder Robertinoida Mikhalevich 1980

Order Robertinida Mikhalevich 1980

Superorder Nonionoida Saidova 1981

Order Elphidiida Saidova 1981

Order Nummulitida Carpenter, Parker & Jones 1862
Order †Orbitoidida Copeland 1956
Order Nonionida Saidova 1981
Superorder Buliminoida Saidova 1981

Order Cassidulinida d’Orbigny 1839

Order Buliminida Saidova 1981
Order Bolivinitida Saidova 1981
Superorder Discorboida Ehrenberg 1838
 Order Chilostomellida Haeckel 1894
Order Discorbida Ehrenberg 1838
Order Glabratellida Mikhalevich 1994
Order Planorbulinida Mikhalevich 1992
Order Rotaliida Lankester 1885
Order Rosalinida Delage & Hérouard 1896
Class Nodosariata Mikhalevich 1992

Subclass Hormosinana Mikhalevich 1992

Order Ammomarginulinida Mikhalevich 2002

Order Nouriida Mikhalevich 1980
Order †Pseudopalmulida Mikhalevich 1992
Order Saccamminida Lankester 1885
Order Hormosinida Mikhalevich 1980
Subclass Nodosariana Mikhalevich 1992

Order †Biseriamminida Mikhalevich 1981

Order Delosinida Revets 1989
Order Lagenida Delage & Hérouard 1896
Order †Palaeotextulariida Hohenegger & Piller 1975
Order Polymorphinida Mikhalevich 1980
Order Vaginulinida Mikhalevich 1993
Order Nodosariida Calkins 1926
Class Spirillinata Mikhalevich 1992

Subclass Ammodiscana Mikhalevich 1980

Order †Plagioraphida Mikhalevich 2003

Order Ammodiscida Mikhalevich 1980 [Pseudoammodiscoida Conil & Lys 1970]
Order Ammovertellinida Mikhalevich 1999
Order Ataxophragmiida Fursenko 1958 [Orbitolinida Ehrenberg 1839]
Subclass Spirillinana Mikhalevich 1992

Superorder †Archaediscoida Pojarkov & Skvortsov 1979

Order †Archaediscida Pojarkov & Skvortsov 1979

Order †Lasiodiscida Mikhalevich 1993
Order †Tetrataxida Mikhalevich 1981
Superorder Involutinoida Hohenegger & Piller 1977

Order †Hottingerellida Mikhalevich 1993

Order Involutinida Hohenegger & Piller 1977
Superorder Spirillinoida Hohenegger & Piller 1975

Order Seabrookiida Mikhalevich 1980

Order Cymbaloporida Mikhaelevich 2013
Order Spirillinida Hohenegger & Piller 1975
Order Patellinida Mikhalevich 1992
Class Miliolata Saidova 1981 (porcelaneous foraminifers)

Subclass Schlumbergerinana Mikhalevich 1992

Order Lituotubida Mikhalevich 1992

Order Loftusiida Kaminski & Mikhalevich 2004
Order Sphaeramminida Mikhalevich & Kaminski 2004
Order Cyclolinida Mikhalevich 1992
 Order Haplophragmiida Loeblich & Tappan 1989
Order Schlumbergerinida Mikhalevich 1980 [Rzehakinida Saidova 1981]
Order Lituolida Lankester 1885
Subclass Miliolana Saidova 1981
Clade Fusulinoids

Order †Ozawainellida Solovieva 1980

Order †Endothyroida Fursenko 1958
Order †Tournayellida Hohenegger & Piller 1973
Order †Fusulinida Fursenko 1958
Order †Neoschwagerinida Minato & Honjo 1966
Order †Schubertellida Skinner 1931
Order †Schwagerinida Solovieva 1985
Order †Staffellida Miklukho-Maklay 1949
Clade Milioloids

Order †Costiferida Mikhalevich 1988

Order Squamulinida Mikhalevich 1988
Order Cornuspirida Jirovec 1953
Order Soritida Schultze 1854 [Orbitolitida Wedekind 1937]
Order Nubeculariida Jones 1875
Order Miliolida Delage & Hérouard 1896

Biology
Modern Foraminifera are primarily marine organisms, but living
individuals have been found in brackish, freshwater[26] and even
terrestrial habitats.[3] The majority of the species are benthic, and a
further 40 morphospecies are planktonic.[27] This count may,
however, represent only a fraction of actual diversity, since many
genetically distinct species may be morphologically
indistinguishable. [28]
Schematic diagram of a live
A number of forams have unicellular algae as endosymbionts, from multilocular foraminifera. 1 -
diverse lineages such as the green algae, red algae, golden algae, endoplasm, 2-ectoplasm, 3-chamber,
diatoms, and dinoflagellates.[27] These mixotrophic foraminifers are 4-pores, 5-foramen, 6-food vacuole,
particularly common in nutrient-poor oceanic waters.[29] Some forams 7-nucleus, 8-mitochondria, 9-
are kleptoplastic, retaining chloroplasts from ingested algae to conduct granureticulose pseudopodia, 10-
granules, 11- primary aperture, 12-
photosynthesis.[30]
food particle, 13-Golgi apparatus, 14-
ribosomes.
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.[26] The pseudopods
are used for locomotion, anchoring, excretion, test construction and in capturing food, which consists of small
organisms such as diatoms or bacteria.[27][31] Certain foraminifera prey upon small animals such as copepods
or cumaceans; some forams even predate upon other forams, drilling holes into the tests of their prey.[32]

Certain benthic foraminifera have been found to be capable of surviving anoxic conditions for over 24 hours,
indicating that they are capable of selective anaerobic respiration. This is interpreted as an adaptation to survive
changing oxygenic conditions near the sediment-water interface.[33]
Reproduction
The generalized foraminiferal life-cycle involves an alternation between haploid and diploid generations,
although they are mostly similar in form.[17][34] The haploid or gamont initially has a single nucleus, and
divides to produce numerous gametes, which typically have two flagella. The diploid or agamont is
multinucleate, and after meiosis divides to produce new gamonts. Multiple rounds of asexual reproduction
between sexual generations are not uncommon in benthic forms.[26]

Foraminifera exhibit morphological

dimorphism associated with their
reproductive cycle. The gamont, or sexually
reproducing haploid form, is megalospheric
—that is, its proloculus, or first chamber, is
proportionally large. The gamont is also
known as the A form. Gamonts, despite
having typically larger proloculi, also
generally have smaller overall test diameter
than do agamonts.

After reaching maturity, the gamont divides

via mitosis to produce thousands of gametes Diagram of a typical foraminiferan life cycle, showing
which are also haploid. These gametes all characteristic alternation of generations.
have a full set of organelles, and are expelled
from the test into the environment leaving
the test undamaged. Gametes are not differentiated into sperm and egg, and any two gametes from a species
can generally fertilize each other.

When two gametes combine, they create a diploid, multi-nucleated

cell known as the agamont, or B form. In contrast to the gamont, the
agamont is microspheric, with a proportionally small first chamber but
typically larger overall diameter with more chambers. The agamont is
the asexual reproduction phase of the foraminifera; upon reaching
adulthood, the protoplasm entirely vacates the test and divides its
cytoplasm meiotically via multiple fission to form a number of haploid
offspring. These offspring then begin to form their megalospheric first
Morphs present in the foram life chamber before dispersing.
cycle—the megalosphere and the
microsphere. The name derives from In some cases the haploid young may mature into a megalospheric
the size of the proloculus, or first form which then reproduces asexually to produce another
chamber, and as such the megalospheric, haploid offspring. In this case, the first megalospheric
microsphere has a larger overall form is referred to as the schizont or A1 form, while the second is
size. referred to as the gamont or A2 form.

Maturation and reproduction occur more slowly in cooler and deeper

water; these conditions also cause forams to grow larger. A forms always seem to be much more numerous
than are B forms, likely due to the reduced likelihood of two gametes encountering one another and
successfully combining.[35][31]

Variations in reproductive mode

There is a high degree of degree of diversity in reproductive strategies in different foraminiferal groups.
In unilocular species, the A form and B form are still present. As in the
microspheric morph of multilocular forams, the asexually reproducing
B form is larger than the sexually reproducing A form.

Forams in the family Spirillinidae have amoeboid gametes rather than

flagellated. Other aspects of reproduction in this group are generally
similar to that of other groups of forams.

The calcareous spirillinid Patellina corrugata has a slightly different

reproductive strategy than most other foraminifera. The asexually Fossil nummulitid foraminiferans
reproducing B form produces a cyst that surrounds the entire cell; it showing microspheric (larger) and
then divides within this cyst and the juvenile cells cannibalise the megalospheric individuals (smaller);
calcite of the parent's test to form the first chamber of their own test. Eocene of the United Arab Emirates;
These A forms, upon maturity, gather into groups of up to 9 scale in mm
individuals; they then form a protective cyst around the whole group.
Gametogenesis occurs within this cyst, producing very low numbers
of gametes. The B form larvae are produced inside of the cyst; any nuclei that are not bound into cells are
consumed as food for the developing larvae. Patellina in A form is reportedly dioecious, with sexes referred to
as the "plus" and "minus"; these sexes differ in number of nuclei, with the "plus" form having three nuclei and
the "minus" form having four nuclei. The B form is again larger than the A form.[31][35][32]

Tests
Foraminiferal tests serve to protect the organism within. Owing to
their generally hard and durable construction (compared to other
protists), the tests of foraminifera are a major source of scientific
knowledge about the group. Openings in the test that allow the
cytoplasm to extend outside are called apertures.[36] The primary
aperture, leading to the exterior, take many different shapes in
different species, including but not limited to rounded, crescent-
shaped, slit-shaped, hooded, radiate (star-shaped), dendritic
(branching). Some foraminifera have "toothed", flanged, or lipped
primary apertures. There may be only one primary aperture or
multiple; when multiple are present, they may be clustered or
Foraminiferan tests (ventral view)
equatorial. In addition to the primary aperture, many foraminifera
have supplemental apertures. These may form as relict apertures (past
primary apertures from an earlier growth stage) or as unique
structures.

Test shape is highly variable among different foraminifera; they may be single-chambered (unilocular) or
multi-chambered (multilocular). In multilocular forms, new chambers are added as the organism grows. A
wide variety of test morphologies is found in both unilocular and multilocular forms, including spiraled, serial,
and milioline, among others.[31]

Many foraminifera exhibit dimorphism in their tests, with megalospheric and microspheric individuals. These
names should not be taken as referring to the size of the full organism; rather, they refer to the size of the first
chamber, or proloculus.

Unlike other shell-secreting organisms, such as molluscs or corals, the tests of foraminifera are located inside
the cell membrane, within the protoplasm. The organelles of the cell are located within the compartment(s) of
the test, and the hole(s) of the test allow the transfer of material from the pseudopodia to the internal cell and
back.[31]
Tests as fossils are known from as far back as the Ediacaran
period,[37] 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.[38] It
is estimated that reef Foraminifera generate about 43 million tons of
calcium carbonate per year.[39]

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.[40]

Test composition

The form and composition of their tests are the primary means by
which forams are identified and classified. Most secrete calcareous
tests, composed of calcium carbonate.[26] Calcareous tests may be
composed of either aragonite or calcite depending on species; among A variety of calcareous foram test
those with calcite tests, the test may contain either a high or low morphologies
fraction of magnesium substitution.[12] The test contains an organic
matrix, which can sometimes be recovered from fossil samples.[12]

Some studies suggest a high amount of homoplasy in foraminifera, and that neither agglutinated nor calcareous
foraminifera form monophyletic groupings.[16]

Soft tests

In some forams, the tests may be composed of organic material, typically the protein tectin. Tectin walls may
have sediment particles loosely adhered onto the surface.[31] The foram Reticulomyxa entirely lacks a test,
having only a membranous cell wall.[41] Organic-walled forams have traditionally been grouped as the
"allogromiids"; however, genetic studies have found that this does not make up a natural group.[16]

Agglutinated tests

Other forams have tests made from small pieces of sediment cemented together (agglutinated) by either
proteins (possibly collagen-related), calcium carbonate, or Iron (III) oxide.[31][42] In the past these forms were
grouped together as the single-chambered "astrorhizids" and the multi-chambered textulariids. However, recent
genetic studies suggest that "astrorhizids" do not make up a natural grouping, instead forming a broad base of
the foram tree.[16]

Agglutinating foraminifera may be selective regarding what particles they incorporate into their shells. Some
species prefer certain sizes and types of rock particles; other species are preferential towards certain biological
materials. Certain species of foraminifera are known to have preferentially agglutinated coccoliths to form their
tests; others preferentially utilise echinoderm plates, diatoms, or even other foraminiferans' tests.[43]

The foraminifera Spiculosiphon preferentially agglutinates silica sponge spicules using an organic cement; it
shows strong selectivity also towards shape, utilising elongated spicules on its "stalk" and shortened ones on
its "bulb". It is thought to use the spicules as both a means of elevating itself off the seabed as well as to
lengthen the reach of its pseudopodia to capture prey.[42]

The agglutinated tests of xenophyophores are the largest of any

foraminifera, reaching up to 20cm in diameter. The name
"xenophyophore", meaning "bearer of foreign bodies", refers to this
agglutinating habit. Xenophyophores selectively uptake sediment
grains between 63 and 500µm, avoiding larger pebbles and finer silts;
type of sediment seems to be a strong factor in which particles are
agglutinated, as particle type preferentially includes sulfides, oxides,
volcanic glass, and especially tests of smaller foraminifera.
Xenophyophores 1.5cm in diameter have been recorded completely
naked, with no test whatsoever.[44] Xenophyophores create the largest
agglutinated tests of any
foraminifera.
Calcareous tests

Of those foraminifera with calcareous tests, several different structures of calcite crystals are found.

Porcelaneous walls are found in the Miliolida. These consist of high-

magnesium calcite organized with an ordered outer and inner calcite
lining (the "extrados" and "intrados", respectively) and randomly
oriented needle-shaped calcite crystals forming a thick center layer
(the "porcelain"). An organic inner lining is also present. The external
surface may have a pitted structure, but it is not perforated by
holes.[45][46]

A "monocrystalline" test
structure has traditionally
The miliolid foraminiferan
been described for the
Quinqueloculina from the North Sea
Spirillinida. However, these
tests remain poorly
understood and poorly
described. Some supposed "monocrystalline" spirillinids have been
found to actually have tests consisting of a mosaic of very small
crystals when observed with scanning electron microscope. SEM SEM photomicrograph of Patellina
observation of Patellina sp. suggests that a truly monocrystalline test sp., showing cleavage of
may indeed be present, with apparent cleavage faces.[47] monocrystalline test

Lagenid tests consist of "fibre bundles" that can reach tens of

micrometres long; each "bundle" is formed from a single calcite crystal, is triangular in cross-section, and has a
pore in the centre (thought to be an artefact of test deposition). There is also an internal organic layer, attached
to the "cone" structure of the fibre bundles. As the crystalline structure varies significantly from that of other
calcareous foraminifera, it is thought to represent a separate evolution of the calcareous test. The exact
mineralisation process of lagenids remains unclear.[48]

Rotaliid tests are described as "hyaline". They are formed from low-to-high-magnesium calcite "nanograins"
positioned with their C-axes perpendicular to the external surface of the test. The test wall is characteristically
bilamellar (two-layered) and perforated throughout with small pores. The outer calcite layer of the test wall is
referred to as the "outer lamina" while the inner calcite layer is referred to as the "inner lining"; this should not
be confused with the organic inner lining beneath the test. Sandwiched between the outer lamina and the inner
lining is the "median layer", a protein layer that separates the two. The median layer is quite variable;
depending on the species it may be well-defined while in others it is not sharply delineated. Some genera may
contain sediment particles within the median layer.[31][49][48]
The Carterinids, including the genera Carterina and Zaninettia, have
a unique crystalline structure of the test which long complicated their
classification. The test in this genus consists of spicules of low-
magnesium calcite, bound together with an organic matrix and
containing "blebs" of organic matter; this led some researchers to
conclude that the test must be agglutinated. However, life studies have
failed to find agglutination, and in fact the genus has been discovered
on artificial substrate where sediment particles do not accumulate.[50]
A 2014 genetic study found carterinids to be an independent lineage
within the Globothalamea, and supported the idea of the spicules
SEM photomicrograph of a lagenid
being secreted as spicule shape differed consistently between
test wall, showing fibre bundle
specimens of Carterina and Zaninettia collected from the same
structures. Internal organic layer can
locality (ovoid in Carterina, rounded-rectangular in Zaninettia).[51] be seen to the far right of the image.

The now-extinct Fusulinids have traditionally been considered unique

in having tests of homogenous microgranular crystals with no
preferred orientation and almost no cement. However, a 2017 study found that the supposed microgranular
structure was actually the result of diagenetic alteration of the fossils, and that unaltered fusulinid tests instead
had a hyaline structure. This suggests that the group is affiliated with the Globothalamea.[47]

Robertinids have aragonitic tests with perforations; these are similar to the tests of rotaliids in that they are
formed from nanograins, however, they differ in composition and in having well-organised columnar domains.
As the earliest planktonic forams had aragonitic tests, it has been suggested that this may represent a separate
evolution of a planktonic lifestyle within the Robertinida, rather than being close relatives of Globigerinans.[46]

Hyaline aragonitic tests are also present in the Involutinida.[48]

Silica tests

One genus, Miliamellus, has a non-perforated test made of opaline silica.[18]

Test wall construction

When a secreted test is present, walls of foraminiferal

tests may be either nonlamellar or lamellar.

Nonlamellar walls are found in some foraminifera, such

as Carterinida, Spirillinida, and Miliolida. In these forms,
the secretion of a new chamber is not associated with any
further deposition over previous chambers. As such there
is no associated layering of calcite layers on the test.[49]

In foraminifera with lamellar walls, the deposition of a

new chamber is accompanied by the deposition of a layer
Anatomy and types of secreted foraminiferal walls over previously-formed chambers. This layer may cover
all previous chambers, or it may cover only some of
them. These layers are known as secondary lamellae.

Foraminifera with lamellar walls can be further broken down into those with monolamellar walls and those
with bilamellar walls. Monolamellar foraminifera secrete test walls which consist of a single layer, while those
of bilamellar foraminifera are double-layered with an organic "median layer", sometimes containing sediment
particles. In the case of bilamellar foraminifera, the outer layer is referred to as the "outer lamella" whilst the
inner layer is referred to as the "inner lining". Monolamellar forams include the Lagenida, while bilamellar
forms include the Rotaliida (including the planktonic subgroup, the Globigerinina).[49]

Bilamellar test walls can be further divided into those with septal flaps (a layer of test wall covering the
previously-secreted septum) and those lacking septal flaps. Septal flaps are not known to be present in any
foraminifera other than those with bilamellar walls.

The presence of a septal flap is often, though not always, associated with the presence of an interlocular
space. As the name suggests, this is a small space located between chambers; it may be open and form part of
the outer surface of the test, or it may be enclosed to form a void. The layer enclosing the void is formed from
different parts of the lamellae in different genera, suggesting an independent evolution of enclosed interlocular
spaces in order to strengthen the test.[49]

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.[52]

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.[52]

Evolutionary history
Molecular clocks indicate that the crown-group of foraminifera likely evolved during the Neoproterozoic,
between 900 and 650 million years ago; this timing is consistent with Neoproterozoic fossils of the closely
related filose amoebae. As fossils of foraminifera have not been found prior to the very end of the Ediacaran, it
is likely that most of these Proterozoic forms did not have hard-shelled tests.[53][54]

Due to their non-mineralised tests, "allogromiids" have no fossil record.[53]

The mysterious vendozoans of the Ediacaran period have been

suggested to represent fossil xenophyophores.[55] However, the
discovery of diagenetically-altered C27 sterols associated with the
remains of Dickinsonia cast doubt on this identification and suggest it
may instead be an animal.[56] Other researchers have suggested that
the elusive trace fossil Paleodictyon and its relatives may represent a
fossil xenophyophore[57] and noted the similarity of the extant
xenophyophore Occultammina to the fossil;[58] however, modern
The mysterious Paleodictyon has examples of Paleodictyon have not been able to clear up the issue and
been interpreted as a fossil the trace may alternately represent a burrow or a glass sponge.[59]
xenophyophore but this remains Supporting this notion is the similar habitat of living xenophyophores
controversial. to the inferred habitat of fossil graphoglyptids; however, the large size
and regularity of many graphoglyptids as well as the apparent absence
of xenophyae in their fossils casts doubt on the possibility.[58] As of
2017 no definite xenophyophore fossils have been found.[60]
Test-bearing foraminifera have an excellent fossil record throughout the Phanerozoic eon. The earliest known
definite foraminifera appear in the fossil record towards the very end of the Ediacaran; these forms all have
agglutinated tests and are unilocular. These include forms like Platysolenites and Spirosolenites.[61][37]

Single-chambered foraminifera continued to diversity throughout the Cambrian. Some commonly encountered
forms include Ammodiscus, Glomospira, Psammosphera, and Turritellella; these species are all agglutinated.
They make up part of the Ammodiscina, a lineage of spirillinids that still contains modern forms.[62][16] Later
spirillinids would evolve multilocularity and calcitic tests, with the first such forms appearing during the
Triassic; the group saw little effects on diversity due to the K-Pg extinction.[63]

The earliest multi-chambered foraminifera are agglutinated species, and appear in the fossil record during the
middle Cambrian period. Due to their poor preservation they cannot be positively assigned to any major foram
group.[62]

The earliest known calcareous-walled foraminifera are the

Fusulinids, which appear in the fossil record during the
Llandoverian epoch of the early Silurian. The earliest of these
were microscopic, planispirally coiled, and evolute; later forms
evolved a diversity of shapes including lenticular, globular, and
perhaps most famously, elongated rice-shaped forms. Later
species of fusulinids grew to much larger size, with some forms
reaching 5 cm in length; reportedly, some specimens reach up to
14 cm in length, making them among the largest foraminifera Cutaway view of a Fusulinid
extant or extinct. Fusulinids are the earliest lineage of foraminifera
thought to have evolved symbiosis with photosynthetic
organisms. Fossils of fusulinids have been found on all continents except Antarctica; they reached their
greatest diversity during the Visean epoch of the Carboniferous. The group then gradually declined in diversity
until finally going extinct during the Permo-Triassic extinction event.[31][63][64]

During the Tournaisian epoch of the Carboniferous, Miliolid foraminifera first appeared in the fossil record,
having diverged from the spirillinids within the Tubothalamea. Miliolids suffered about 50% casualties during
both the Permo-Triassic and K-Pg extinctions but survived to the present day. Some fossil miliolids reached up
to 2 cm in diameter.[63]

The earliest known Lagenid fossils appear during the Moscovian

epoch of the Carboniferous. Seeing little effect due to the Permo-
Triassic or K-Pg extinctions, the group diversified through time.
Secondarily unilocular taxa evolved during the Jurassic and
Cretaceous.

The earliest Involutinid fossils appear during the Permian; the lineage
diversified throughout the Mesozoic of Eurasia before apparently
vanishing from the fossil record following the Cenomanian-Turonian
Ocean Anoxic Event. The extant group planispirillinidae has been
referred to the involutinida, but this remains the subject of
debate.[65][63]

A fossil test from a planktonic The Robertinida first appear in the fossil record during the Anisian
globigerininan foraminifera. epoch of the Triassic. The group remained at low diversity throughout
its fossil history; all living representatives belong to the Robertinidae,
which first appeared during the Paleocene.[63]
The first definite Rotaliid fossils do not appear in the fossil record until the Pliensbachian epoch of the Jurassic,
following the Triassic-Jurassic event.[66] Diversity of the group remained low until the aftermath of the
Cenomanian-Turonian event, after which the group saw a rapid diversification. Of this group, the planktonic
Globigerinina first appear in the aftermath of the Toarcian Turnover; the group saw heavy losses during both
the K-Pg extinction and the Eocene-Oligocene extinction, but remains extant and diverse to this day.[63]

Paleontological applications
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.[67] 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.[67]
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 Neoflabellina reticulata from chalk of
Rügen, Northeastern Germany.
specimens[68]
Length:1.2 mm, Age: Upper lower
Maastrichtian
Because certain types of foraminifera are found only in certain
environments, their fossils can be used to figure out the kind of
environment under which ancient marine sediments were deposited;
conditions such as salinity, depth, oxygenic conditions, and light
conditions can be determined from the different habitat preferences of
various species of forams. This allows workers to track changing
climates and environmental conditions over time by aggregating
information about the foraminifera present.[69]

In other cases, the relative proportion of planktonic to benthic

foraminifera fossils found in a rock can be used as a proxy for the
Thin section of a peneroplid
depth of a given locality when the rocks were being deposited.[70]
foraminiferan from Holocene lagoonal
sediment in Rice Bay, San Salvador
Foraminifera have significant application in the field of
Island, Bahamas. Scale bar 100
biostratigraphy. Due to their small size and hard shells, foraminifera
micrometres
may be preserved in great abundance and with high quality of
preservation; due to their complex morphology, individual species are
easily recognizable. Foraminifera species in the fossil record have
limited ranges between the species' first evolution and their disappearance; stratigraphers have worked out the
successive changes in foram assemblages throughout much of the Phanerozoic. As such, the assemblage of
foraminifera within a given locality can be analyzed and compared to known dates of appearance and
disappearance in order to narrow down the age of the rocks. This allows paleontologists to interpret the age of
sedimentary rocks when radiometric dating is not applicable.[71] This application of foraminifera was
discovered by Alva C. Ellisor in 1920.[72]
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;[73] see δ18O and δ13C. The
concentration of trace elements, like magnesium (Mg),[74] lithium (Li)[75] and boron (B),[76] 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.

Modern uses
The oil industry relies heavily on microfossils such as forams to find potential hydrocarbon deposits.[77]

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.[78] Agglutinated fossil foraminifera buried deeply in
Ammonia beccarii, a benthic foram
sedimentary basins can be used to estimate thermal maturity, which is
from the North Sea.
a key factor for petroleum generation. The Foraminiferal Colouration
Index[79] (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".[80]

Gallery Foraminifera Baculogypsina

sphaerulata of Hatoma Island,
Japan. Field width 5.22 mm
Foraminifera of Pag Foraminifera of Pag Foraminifera of Pag Foraminifera of Pag
Island, Adriatic Sea Island, Adriatic Sea Island, Adriatic Sea Island, Adriatic Sea
-60 m, field width -60 m, field width -60 m, field width -60 m, field width
5.5 mm 5.5 mm 5.5 mm 5.5 mm

Foraminifera of Foraminifera of Foraminifera of Foraminifera in

Indian Ocean, south- Indian Ocean, south- Indian Ocean, south- Ngapali, Myanmar,
eastern coast of eastern coast of eastern coast of field width 5.22 mm
Bali, field width Bali, field width Bali, field width
5.5 mm 5.5 mm 5.5 mm

Foraminifera
Heterostegina
depressa, field width
4.4 mm

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External links
General information

The University of California Museum of Paleontology ([Link]

l) website has an Introduction to the Foraminifera ([Link]
[Link])
Researchers at the University of South Florida developed a system using Foraminifera for
monitoring coral reef environments ([Link]
[Link])
University College London's micropaleontology site ([Link]
[Link]) has an overview of Foraminifera, including many high-quality SEMs
Illustrated glossary of terms used in foraminiferal research ([Link]
06_M02/[Link]) is the Lukas Hottinger's glossary published in the OA e-journal "Carnets
de Géologie — Notebooks on Geology" ([Link]
 Information on Foraminifera ([Link]
[Link]/[Link]) Martin Langer's Micropaleontology Page
Benthic Foraminifera information ([Link] from the
2005 Urbino Summer School of Paleoclimatology

Online flip-books

Illustrated glossary of terms used in foraminiferal research ([Link]

34b9de489f0c3c) by Lukas Hottinger (alternative version of the one published in "Carnets de
Géologie — Notebooks on Geology" ([Link]

Resources

pforams@mikrotax ([Link] - an online database detailing the

taxonomy of planktonic foraminifera
The star*sand project ([Link]
[Link]) (part of micro*scope ([Link]
[Link]/mv/[Link]?pagetitle=index)) is a cooperative database of information about
Foraminifera
3D models ([Link] of forams, generated by X-ray
tomography
CHRONOS ([Link] has
several Foraminifera resources ([Link]
[Link]/gridsphere/gridsphere?cid=res_foram), including a taxon search page ([Link]
[Link]/web/20060512035435/[Link]
b) and a micro-paleo section ([Link]
[Link]/gridsphere/gridsphere?cid=micropaleo) NB Most of this content is now included in the
pforams@mikrotax website
eForams ([Link] is a web site focused on Foraminifera and modeling of
foraminiferal shells
Foraminifera Gallery ([Link] Illustrated catalog of recent and fossil
Foraminifera by genus and locality
"Foraminifera" ([Link]
9178&lvl=1). NCBI Taxonomy Browser. 29178.

Retrieved from "[Link]

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