Foraminifera - Wikipedia

7/3/2021 Foraminifera - Wikipedia
Foraminifera
Foraminifera (/fəˌræməˈnɪfərə/; Latin for "hole bearers"; informally
called "forams") are single-celled organisms, members of a phylum or Foraminifera
class of amoeboid protists characterized by streaming granular ectoplasm Temporal range: 542–0 Ma[1]
for catching food and other uses; and commonly an external shell (called
PreꞒ Ꞓ O S D C P T J K PgN
a "test") of diverse forms and materials. Tests of chitin (found in some Latest Ediacaran–Recent
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),[2] while a smaller
number float in the water column at various depths (i.e., are planktonic),
which belong to the suborder Globigerinina.[3] Fewer are known from
freshwater[4] or brackish[5] conditions, and some very few (nonaquatic)
soil species have been identified through molecular analysis of small
subunit ribosomal DNA.[6][7]
Foraminifera typically produce a test, or shell, which can have either one Live Ammonia tepida (Rotaliida)
or multiple chambers, some becoming quite elaborate in structure.[8]
Scientific classification
These shells are commonly made of calcium carbonate (CaCO
3) or
agglutinated sediment particles. Over 50,000 species are recognized, both Domain: Eukaryota
living (6,700 - 10,000)[9][10] and fossil (40,000).[11][12] They are usually (unranked): SAR
less than 1 mm in size, but some are much larger, the largest species
reaching up to 20 cm.[13] (unranked): Rhizaria
Phylum: Retaria
In modern scientific English, the term foraminifera is both singular and
plural (irrespective of the word's Latin derivation), and is used to Subphylum: Foraminifera
describe one or more specimens or taxa: its usage as singular or plural d'Orbigny, 1826
must be determined from context. Foraminifera is frequently used
informally to describe the group, and in these cases is generally Subdivisions
lowercase.[14]
"Monothalamea"
"Allogromiida"
Contents
"Astrorhizida"
History of study
Xenophyophorea
Taxonomy
Reticulomyxa
Anatomy
Ecology Tubothalamea
Reproduction
Variations in reproductive mode Miliolida
Tests Spirillinida
Composition Silicoloculinida
Soft
Agglutinated Globothalamea
https://en.wikipedia.org/wiki/Foraminifera 1/27
Calcareous Textulariida
Silicate
Rotaliida
Test wall construction
Evolutionary history Globigerinida
Paleontological applications Carterinida

Modern uses Robertinida
Gallery Fusulinida? — extinct
References
incertae sedis
External links
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.[15]
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[...][16]
Antonie van Leeuwenhoek described and illustrated foraminiferal tests in 1700, describing them as minute cockles;
his illustration is recognizable as being Elphidium.[17] 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.[18] Spengler also noted that
the septa of foraminifera arced the opposite way from those of nautili and that they lacked a nerve tube.[19]
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.[20]
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.[14]
The protozoan nature of foraminifera was first recognized by Dujardin in 1835.[18] 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.[17]
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.[17] 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 groups.[21] Earliest known illustration of a
foraminifera shell, published by
Taxonomy Robert Hooke in his 1665 book
Micrographia.
The taxonomic position of the Foraminifera has varied since Schultze in

1854,[22] who referred to as an order, Foraminiferida. Loeblich
(1987) and Tappan (1992) reranked Foraminifera as a class[23]  
"Monothalamids" (paraphyletic)
as it is now commonly regarded.  
The Foraminifera have typically been included in the  

  Lagenida
Protozoa,[24][25][26] or in the similar Protoctista or Protist
kingdom.[27][28] Compelling evidence, based primarily on  
molecular phylogenetics, exists for their belonging to a major "Monothalamids"
 
group within the Protozoa known as the Rhizaria.[24] Prior to  
the recognition of evolutionary relationships among the     
members of the Rhizaria, the Foraminifera were generally     Miliolida
Tubothalamea
grouped with other amoeboids as phylum Rhizopodea (or  
Sarcodina) in the class Granuloreticulosa.  
Spirillinida
 
The Rhizaria are problematic, as they are often called a  
"supergroup", rather than using an established taxonomic rank     "Monothalamids"
such as phylum. Cavalier-Smith defines the Rhizaria as an    
infra-kingdom within the kingdom Protozoa.[24]  
 
  Xenophyophorea
Some taxonomies put the Foraminifera in a phylum of their
own, putting them on par with the amoeboid Sarcodina in   "Textulariida"
which they had been placed.  
    (paraphyletic)
Although as yet unsupported by morphological correlates, Globothalamea
molecular data strongly suggest the Foraminifera are closely  
  Robertinida
related to the Cercozoa and Radiolaria, both of which also  
include amoeboids with complex shells; these three groups  
make up the Rhizaria.[25] However, the exact relationships of   Rotaliida
the forams to the other groups and to one another are still not
entirely clear. Foraminifera are closely related to testate Phylogeny of Foraminifera following Pawlowski et
amoebae.[29] al. 2013.[21] The monothalamid orders
Astrorhizida and Allogromiida are both

paraphyletic.
Taxonomy from Mikhalevich 2013[30]

* 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
Anatomy
The most striking aspect of most foraminifera are their hard shells, or tests.
These may consist of one of multiple chambers, and may be composed of
protein, sediment particles, calcite, aragonite, or (in one case) silica.[23]
Some foraminifera lack tests entirely.[31] 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 Schematic diagram of a live
back.[32] multilocular foraminifera. 1 -
endoplasm, 2-ectoplasm, 3-
The foraminiferal cell is divided into granular endoplasm and transparent chamber, 4-pores, 5-foramen, 6-
ectoplasm from which a pseudopodial net may emerge through a single food vacuole, 7-nucleus, 8-
opening or through many perforations in the test. Individual pseudopods mitochondria, 9-granureticulose
characteristically have small granules streaming in both directions.[33] pseudopodia, 10-granules, 11-
Foraminifera are unique in having granuloreticulose pseudopodia; that is, primary aperture, 12-food particle,
13-Golgi apparatus, 14-ribosomes.
their pseudopodia appear granular under the microscope; these
pseudopodia are often elongate and may split and rejoin each other. These
can be extended and retracted to suit the needs of the cell. The pseudopods
are used for locomotion, anchoring, excretion, test construction and in capturing food, which consists of small
organisms such as diatoms or bacteria.[34][32]
Aside from the tests, foraminiferal cells are supported by a cytoskeleton of microtubules, which are loosely
arranged without the structure seen in other amoeboids. Forams have evolved special cellular mechanisms to
quickly assemble and disassemble microtubules, allowing for the rapid formation and retraction of elongated
pseudopodia.[23]
In the gamont (sexual form), foraminifera generally have only a single nucleus, while the agamont (asexual form)
tends to have multiple nuclei. In at least some species the nuclei are dimorphic, with the somatic nuclei containing
three times as much protein and RNA than the generative nuclei. However, nuclear anatomy seems to be highly
diverse.[35] The nuclei are not necessarily confined to one chamber in multi-chambered species. Nuclei can be
spherical or have many lobes. Nuclei are typically 30-50 µm in diameter.[36]
Some species of foraminifera have large, empty vacuoles within their cells; the exact purpose of these is unclear,
but they have been suggested to function as a reservoir of nitrate.[36]
Mitochondria are distributed evenly throughout the cell, though in some species they are concentrated under the
pores and around the external margin of the cell. This has been hypothesised to be an adaptation to low-oxygen
environments.[36]
Several species of xenophyophore have been found to have unusually high concentrations of radioactive isotopes
within their cells, among the highest of any eukaryote. The purpose of this is unknown.[37]
Ecology
Modern Foraminifera are primarily marine organisms, but living individuals have been found in brackish,
freshwater[33] and even terrestrial habitats.[7] The majority of the species are benthic, and a further 40
morphospecies are planktonic.[34] This count may, however, represent only a fraction of actual diversity, since
many genetically distinct species may be morphologically indistinguishable.[38]
Benthic foraminifera are typically found in fine-grained sediments, where they actively move between layers;
however, many species are found on hard rock substrates, attached to seaweeds, or sitting atop the sediment
surface.[23]
The majority of planktonic foraminifera are found in the globigerinina, a lineage within the rotaliida.[21] However,
at least one other extant rotaliid lineage, Neogallitellia, seems to have independently evolved a planktonic
lifestyle.[39][40] Further, it has been suggested that some Jurassic fossil foraminifera may have also independently
evolved a planktonic lifestyle, and may be members of Robertinida.[41]
A number of forams have unicellular algae as endosymbionts, from diverse lineages such as the green algae, red
algae, golden algae, diatoms, and dinoflagellates.[34] These mixotrophic foraminifers are particularly common in
nutrient-poor oceanic waters.[42] Some forams are kleptoplastic, retaining chloroplasts from ingested algae to
conduct photosynthesis.[43]
Most foraminifera are heterotrophic, consuming smaller organisms and organic matter; some smaller species are
specialised feeders on phytodetritus, while others specialise in consuming diatoms. Some benthic forams construct
feeding cysts, using the pseuodopodia to encyst themselves inside of sediment and organic particles.[23] 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.[44] One group, the xenophyophores, has been suggested to farm
bacteria within their tests.[45] Suspension feeding is also common in the group, and at least some species can take
advantage of dissolved organic carbon.[23]
A few foram species are parasitic, infecting sponges, molluscs, corals, or even other foraminifera. Parasitic
strategies vary; some act as ectoparasites, using their pseudopodia to steal food from the host, while others burrow
through the shell or body wall of their host to feed on its soft tissue.[23]
Foraminifera are themselves eaten by a host of larger organisms, including invertebrates, fish, shorebirds, and other
foraminifera. It has been suggested, however, that in some cases predators may be more interested in the calcium
from foram shells than in the organisms themselves. Several aquatic snail species are known to selectively feed
upon foraminifera, often even preferring individual species.[46]
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.[47]
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.[48]
Reproduction
The generalized foraminiferal life-cycle involves an alternation between haploid and diploid generations, although
they are mostly similar in form.[22][49] 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.[33]
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 which are
also haploid. These gametes all have a full set of Diagram of a typical foraminiferan life cycle, showing
organelles, and are expelled from the test into the characteristic alternation of generations.
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
chamber before dispersing.
In some cases the haploid young may mature into a megalospheric form
which then reproduces asexually to produce another megalospheric,
haploid offspring. In this case, the first megalospheric form is referred to as
the schizont or A1 form, while the second is referred to as the gamont or A2
form.
Maturation and reproduction occur

more slowly in cooler and deeper
Morphs present in the foram life water; these conditions also cause
cycle—the megalosphere and the forams to grow larger. A forms
microsphere. The name derives always seem to be much more
from the size of the proloculus, or numerous than are B forms, likely
first chamber, and as such the due to the reduced likelihood of
microsphere has a larger overall two gametes encountering one
size. another and successfully
combining. [50][32] Fossil nummulitid foraminiferans
showing microspheric (larger) and
megalospheric individuals (smaller);
Variations in reproductive mode Eocene of the United Arab
Emirates; scale in mm
There is a high 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 reproducing B form produces a cyst that surrounds the entire cell; it then divides
within this cyst and the juvenile cells cannibalise the calcite of the parent's test to form the first chamber of their
own test. These A forms, upon maturity, gather into groups of up to nine 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.[32][50][44]
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.[51] 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 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.[32]
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.
Foraminiferan tests (ventral view) Tests as fossils are known from as

far back as the Ediacaran period,[52]
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.[53] It is estimated that reef Foraminifera generate about 43
million tons of calcium carbonate per year.[54]
Genetic studies have identified the naked amoeba Reticulomyxa and the
peculiar xenophyophores as foraminiferans without tests. A few other The miliolid foraminiferan
amoeboids produce reticulose pseudopods, and were formerly classified Quinqueloculina from the North Sea
with the forams as the Granuloreticulosa, but this is no longer considered a
natural group, and most are now placed among the Cercozoa.[55]
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.[33] Calcareous tests may be composed of either aragonite
or calcite depending on species; among those with calcite tests, the test may contain either a high or low fraction of
magnesium substitution.[17] The test contains an organic matrix, which can sometimes be recovered from fossil
samples.[17]
Some studies suggest a high amount of homoplasy in foraminifera, and that neither agglutinated nor calcareous
foraminifera form monophyletic groupings.[21]
Soft
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.[32] The foram Reticulomyxa entirely lacks a test, having only a
membranous cell wall.[31] Organic-walled forams have traditionally been grouped as the "allogromiids"; however,
genetic studies have found that this does not make up a natural group.[21]
Agglutinated
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.[32][56] 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.[21]
Textulariid foraminifera, unlike other living members of the globothalamea, have agglutinated tests; however,
grains in these tests are cemented with a calcite cement. This calcite cement is made up of small (<100 nm)
globular nanograins, similar to in other globothalameans. These tests may also have many pores, another feature
uniting them with the globothalamea.[41]
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.[57]
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 Xenophyophores create the largest
agglutinated tests of any
prey.[56]
foraminifera.
The agglutinated tests of xenophyophores are the largest of any
foraminifera, reaching up to 20 cm in diameter. The name
"xenophyophore", meaning "bearer of foreign bodies", refers to this agglutinating habit. Xenophyophores
selectively uptake sediment grains between 63 and 500 µm, 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 1.5 cm in diameter
have been recorded completely naked, with no test whatsoever.[58]
Calcareous
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
SEM photomicrograph of a miliolid with an ordered outer and inner
test wall, showing the nanogranular calcite lining (the "extrados" and
extrados (e) and needlelike porcelain "intrados", respectively) and
(p) layers.
randomly oriented needle-shaped
calcite crystals forming a thick
center layer (the "porcelain"). An SEM photomicrograph of Patellina
organic inner lining is also present. The external surface may have a pitted sp., showing cleavage of
structure, but it is not perforated by holes. "Cornuspirid" miliolids monocrystalline test
apparently lack any extrados. [59][60][41]
A "monocrystalline" test structure
has traditionally been described for the 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 observation of Patellina sp. suggests that a truly monocrystalline test may indeed be present, with apparent
cleavage faces.[41]
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.[60]
Rotaliid tests are described as

SEM photomicrograph of a lagenid
"hyaline". They are formed from
test wall, showing fibre bundle
low-to-high-magnesium calcite
structures. Internal organic layer can
be seen to the far right of the image.
"nanograins" positioned with their
C-axes perpendicular to the external
surface of the test. Further, these
nanograins can have higher-level structure, such as rows, columns, or
bundles.[41] The test wall is characteristically bilamellar (two-layered) and
perforated throughout with small pores. The outer calcite layer of the test SEM photomicrograph showing
wall is referred to as the "outer lamina" while the inner calcite layer is cross section of a rotaliid wall. Note
referred to as the "inner lining"; this should not be confused with the organic "globular nanograins" and two-
inner lining beneath the test. Sandwiched between the outer lamina and the layered test wall. Arrows point to
inner lining is the "median layer", a protein layer that separates the two. The pores.
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.[32][61][60]
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.[62] A 2014 genetic study found carterinids to
be an independent lineage within the Globothalamea, and supported the
idea of the spicules being secreted as spicule shape differed consistently
SEM photomicrograph of a between specimens of Carterina and Zaninettia collected from the same
carterinid test wall, showing locality (ovoid in Carterina, rounded-rectangular in Zaninettia).[63]
secreted calcite spicules in organic
matrix. 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.[64]
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.[41]
Hyaline aragonitic tests are also present in the Involutinida.[60]
Silicate
One genus, Miliamellus, has a non-perforated test made of opaline silica.[23] It is similar in shape and structure to
the tests of typical miliolids; the test consists of an internal and external organic layer, as well as a middle silica
layer made of elongate rods. This silica layer is further divided into outer, middle, and inner subunits; the outer and
inner subunits each are approximately 0.2μm thick and consist of subparallel sheets of silica rods with their long
axes parallel to the test surface. The middle subunit is approximately 18μm in thickness and consists of a three-
dimensional lattice of silica rods with no organic component in the open space. The ultrastructure differs from that
of miliolids in that the rods are over twice as long and twice as thick on average, in that the rods of Miliamellus are
hollow rather than solid, and of course in having a silica test rather than calcite.[65]
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.[61]
In foraminifera with lamellar walls, the deposition of a new

chamber is accompanied by the deposition of a layer over
Anatomy and types of secreted foraminiferal walls 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 major planktonic subgroup, the Globigerinina).[61]
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.[61]
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.[66][67]
Due to their non-mineralised tests, "allogromiids" have no fossil record.[66]
The mysterious vendozoans of the Ediacaran period have been suggested to

represent fossil xenophyophores.[68] 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.[69] Other researchers have suggested that the elusive trace fossil
Paleodictyon and its relatives may represent a fossil xenophyophore[70] and
noted the similarity of the extant xenophyophore Occultammina to the
fossil;[71] however, modern examples of Paleodictyon have not been able
The mysterious Paleodictyon has to clear up the issue and the trace may alternately represent a burrow or a
been interpreted as a fossil glass sponge.[72] Supporting this notion is the similar habitat of living
xenophyophore but this remains xenophyophores to the inferred habitat of fossil graphoglyptids; however,
controversial. the large size and regularity of many graphoglyptids as well as the apparent
absence of xenophyae in their fossils casts doubt on the possibility.[71] As
of 2017 no definite xenophyophore fossils have been found.[73]
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.[74][52]
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.[75][21] 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.[76]
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.[75]
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 extant or
extinct. Fusulinids are the earliest lineage of foraminifera thought to Cutaway view of a Fusulinid
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.[32][76][77]
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.[76]
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.[78][76]
The Robertinida first appear in the fossil record during the Anisian epoch
of the Triassic. The group remained at low diversity throughout its fossil
A fossil test from a planktonic history; all living representatives belong to the Robertinidae, which first
globigerininan foraminifera. appeared during the Paleocene.[76]
The first definite Rotaliid fossils do not appear in the fossil record until the
Pliensbachian epoch of the Jurassic, following the Triassic-Jurassic event.[79] 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—the first known group of planktonic forams—first appears 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.[76] An additional evolution of planktonic lifestyle occurred in
the Miocene or Pliocene, when the rotaliid Neogallitellia independently evolved a planktonic lifestyle.[39][40]
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.[80] 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.[80] 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[81]
Neoflabellina reticulata from chalk of
Because certain types of foraminifera are found only in certain Rügen, Northeastern Germany.
environments, their fossils can be used to figure out the kind of Length:1.2 mm, Age: Upper lower
Maastrichtian
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.[82]
In other cases, the relative proportion of planktonic to benthic foraminifera

fossils found in a rock can be used as a proxy for the depth of a given
locality when the rocks were being deposited.[83]
Thin section of a peneroplid
Foraminifera have significant application in the field of biostratigraphy. foraminiferan from Holocene
Due to their small size and hard shells, foraminifera may be preserved in lagoonal sediment in Rice Bay, San
great abundance and with high quality of preservation; due to their complex Salvador Island, Bahamas. Scale
morphology, individual species are easily recognizable. Foraminifera bar 100 micrometres
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.[84] This application of foraminifera was discovered
by Alva C. Ellisor in 1920.[85]
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;[86] see δ18O and δ13C. The concentration of trace
elements, like magnesium (Mg),[87] lithium (Li)[88] and boron (B),[89] 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.[90]
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.[91] Agglutinated fossil foraminifera buried deeply in
sedimentary basins can be used to estimate thermal maturity, which is a key
Ammonia beccarii, a benthic foram
factor for petroleum generation. The Foraminiferal Colouration Index[92] from the North Sea.
(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".[93]
Gallery

Foraminifera Baculogypsina
sphaerulata of Hatoma Island,
Japan. Field width 5.22 mm
Foraminifera of Pag Foraminifera of Pag Foraminifera of Pag

Island, Adriatic Sea Island, Adriatic Sea Island, Adriatic Sea
-60 m, field width -60 m, field width -60 m, field width
5.5 mm 5.5 mm 5.5 mm
Foraminifera of Pag Foraminifera of Foraminifera of Foraminifera of

Island, Adriatic Sea Indian Ocean, Indian Ocean, Indian Ocean,
-60 m, field width south-eastern coast south-eastern coast south-eastern coast
5.5 mm of Bali, field width of Bali, field width of Bali, field width
5.5 mm 5.5 mm 5.5 mm
Foraminifera in Foraminifera
Ngapali, Myanmar, Heterostegina
field width 5.22 mm depressa, field width
4.4 mm
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development in SW Hungary: subsidence history based on quantitative paleobathymetry of
foraminifera". International Journal of Earth Sciences. 91 (3): 490–504.
Bibcode:2002IJEaS..91..490B (https://ui.adsabs.harvard.edu/abs/2002IJEaS..91..490B).
doi:10.1007/s005310100226 (https://doi.org/10.1007%2Fs005310100226). ISSN 1437-3262 (https://
www.worldcat.org/issn/1437-3262). S2CID 129296067 (https://api.semanticscholar.org/CorpusID:129
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85. Cushman, Joseph A.; Ellisor, Alva C. (1 January 1945). "The Foraminiferal Fauna of the Anahuac
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External links
General information
The University of California Museum of Paleontology (http://www.ucmp.berkeley.edu/index.html)

website has an Introduction to the Foraminifera (http://www.ucmp.berkeley.edu/foram/foramintro.htm
l)
Researchers at the University of South Florida developed a system using Foraminifera for monitoring
coral reef environments (http://www.marine.usf.edu/reefslab/foramcd/html_files/titlepage.htm)
University College London's micropaleontology site (http://www.ucl.ac.uk/GeolSci/micropal/foram.ht
ml) has an overview of Foraminifera, including many high-quality SEMs
Illustrated glossary of terms used in foraminiferal research (http://paleopolis.rediris.es/cg/CG2006_M
02/index.html) is the Lukas Hottinger's glossary published in the OA e-journal "Carnets de Géologie
— Notebooks on Geology" (http://paleopolis.rediris.es/cg/uk-index.html)
Information on Foraminifera (https://web.archive.org/web/20070810004729/http://www.paleontology.
uni-bonn.de/frame02.htm) Martin Langer's Micropaleontology Page
Benthic Foraminifera information (http://ethomas.web.wesleyan.edu/BFhandout.htm) from the 2005
Urbino Summer School of Paleoclimatology
Online flip-books
Illustrated glossary of terms used in foraminiferal research (http://fr.calameo.com/read/000023034b9

de489f0c3c) by Lukas Hottinger (alternative version of the one published in "Carnets de Géologie —
Notebooks on Geology" (http://paleopolis.rediris.es/cg/uk-index.html))
Resources
pforams@mikrotax (http://www.mikrotax.org/pforams/) - an online database detailing the taxonomy of

planktonic foraminifera
The star*sand project (https://web.archive.org/web/20060620040556/http://www.bowserlab.org/stars
and.html) (part of micro*scope (https://web.archive.org/web/20080609213132/http://starcentral.mbl.e
du/mv/portal.php?pagetitle=index)) is a cooperative database of information about Foraminifera
3D models (http://webdb2.museum.tohoku.ac.jp/e-foram/) of forams, generated by X-ray tomography
CHRONOS (https://web.archive.org/web/19981205122129/http://www.chronos.org/) has several
Foraminifera resources (https://web.archive.org/web/20061115112221/http://portal.chronos.org/grids
phere/gridsphere?cid=res_foram), including a taxon search page (https://web.archive.org/web/20060
512035435/http://portal.chronos.org/gridsphere/gridsphere?cid=res_taxondb) and a micro-paleo
section (https://web.archive.org/web/20120328221949/http://portal.chronos.org/gridsphere/gridspher
e?cid=micropaleo) NB Most of this content is now included in the pforams@mikrotax website
eForams (http://www.eforams.org) is a web site focused on Foraminifera and modeling of
foraminiferal shells
Foraminifera Gallery (http://www.foraminifera.eu) Illustrated catalog of recent and fossil Foraminifera
by genus and locality
"Foraminifera" (https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Info&id=29178&
lvl=1). NCBI Taxonomy Browser. 29178.
Retrieved from "https://en.wikipedia.org/w/index.php?title=Foraminifera&oldid=1025209710"
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7/3/2021 Foraminifera - Wikipedia

Paleontological applications Carterinida

This overall scheme of classification would remain until Cushman's work

Cushman's scheme nevertheless remained the dominant scheme of

The taxonomic position of the Foraminifera has varied since Schultze in

The Foraminifera have typically been included in the

Astrorhizida and Allogromiida are both

Taxonomy from Mikhalevich 2013[30]

Order Stannomida Tendal 1972

Subclass Lagynana Mikhalevich 1980

Order Ammoscalariida Mikhalevich 1980

Order Astrorhizida Lankester 1885

Subclass Globigerinana Mikhalevich 1980

Order Cassigerinellida Mikhalevich 2013

Order Nautiloculinida Mikhalevich 2003

Superorder Robertinoida Mikhalevich 1980

Order Robertinida Mikhalevich 1980

Order Elphidiida Saidova 1981

Order Cassidulinida d’Orbigny 1839

Order Bolivinitida Saidova 1981

Order Chilostomellida Haeckel 1894

Subclass Hormosinana Mikhalevich 1992

Order Ammomarginulinida Mikhalevich 2002

Order †Biseriamminida Mikhalevich 1981

Subclass Ammodiscana Mikhalevich 1980

Order †Plagioraphida Mikhalevich 2003

Superorder †Archaediscoida Pojarkov & Skvortsov 1979

Order †Archaediscida Pojarkov & Skvortsov 1979

Order †Hottingerellida Mikhalevich 1993

Order Seabrookiida Mikhalevich 1980

Subclass Schlumbergerinana Mikhalevich 1992

Order Lituotubida Mikhalevich 1992

Order Loftusiida Kaminski & Mikhalevich 2004

Order †Ozawainellida Solovieva 1980

Order †Costiferida Mikhalevich 1988

Foraminifera exhibit morphological dimorphism

After reaching maturity, the gamont divides via

Maturation and reproduction occur

Test shape is highly variable among different foraminifera; they may be

Many foraminifera exhibit dimorphism in their tests, with megalospheric

Foraminiferan tests (ventral view) Tests as fossils are known from as

Agglutinating foraminifera may be selective regarding what particles they

The foraminifera Spiculosiphon preferentially agglutinates silica sponge

Of those foraminifera with calcareous tests, several different structures of

Porcelaneous walls are found in

Rotaliid tests are described as

The Carterinids, including the genera Carterina and Zaninettia, have a

Hyaline aragonitic tests are also present in the Involutinida.[60]

Test wall construction

When a secreted test is present, walls of foraminiferal tests

Nonlamellar walls are found in some foraminifera, such as

In foraminifera with lamellar walls, the deposition of a new

Due to their non-mineralised tests, "allogromiids" have no fossil record.[66]

The mysterious vendozoans of the Ediacaran period have been suggested to

The earliest known calcareous-walled foraminifera are the Fusulinids,

can be determined from the different habitat preferences of various species

In other cases, the relative proportion of planktonic to benthic foraminifera

Foraminifera have many uses in petroleum exploration and are used

Foraminifera can also be used in archaeology in the provenancing of some

Foraminifera of Pag Foraminifera of Pag Foraminifera of Pag

Foraminifera of Pag Foraminifera of Foraminifera of Foraminifera of

24. Cavalier-Smith, T (2004). "Only Six Kingdoms of Life" (http://www.cladocera.de/protozoa/cavalier-smi

The University of California Museum of Paleontology (http://www.ucmp.berkeley.edu/index.html)

Illustrated glossary of terms used in foraminiferal research (http://fr.calameo.com/read/000023034b9

The Foraminifera have typically been included in the  