Foraminifera ( ; Latin for "hole bearers"; informally called "forams") are single-celled organisms, members of a phylum or class of Rhizarian protists characterized by streaming granular ectoplasm for catching food and other uses; and commonly an external shell called a test of diverse forms and materials. Tests of chitin (found in some simple genera, and Textularia in particular) are believed to be the most primitive type. Most foraminifera are marine, the majority of which live on or within the seafloor sediment (i.e., are benthic, with different sized species playing a role within the macrobenthos, meiobenthos, and microbenthos), while a smaller number float in the water column at various depths (i.e., are planktonic), which belong to the suborder Globigerinina. 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.
Foraminifera typically produce a foraminifera test, or shell, which can have either one or multiple chambers, some becoming quite elaborate in structure. These shells are commonly made of calcium carbonate () or agglutinated sediment particles. Over 50,000 species are recognized, both living (6,700–10,000)Ald, S.M. et al. (2007) Diversity, Nomenclature, and Taxonomy of Protists, Syst. Biol. 56(4), 684–689, DOI: 10.1080/10635150701494127. and fossil (40,000).Pawlowski, J., Lejzerowicz, F., & Esling, P. (2014). Next-generation environmental diversity surveys of foraminifera: preparing the future. The Biological Bulletin, 227(2), 93-106. They are usually less than 1 mm in size, but some are much larger, the largest species reaching up to 20 cm.
In modern scientific English, the term foraminifera is both singular and plural (irrespective of the word's Latin derivation), and is used to describe one or more specimens or taxa: its usage as singular or plural must be determined from context. Foraminifera is frequently used informally to describe the group, and in these cases is generally lowercase.
History of study The earliest known reference to foraminifera comes from Strabo, who in the 1st century BCE noted them as making up the rock that forms the Great Pyramid of Giza and suggested that they were the remains of lentils left by the workers who built the pyramids.Strabo, Geographica, XVII.1.34. These are today recognized as representatives of the genus Nummulites.
Robert Hooke observed a foraminifera under the microscope, as described and illustrated in his 1665 book Micrographia:
Antonie van Leeuwenhoek described and illustrated foraminiferal tests in 1700, describing them as minute cockles; his illustration is recognizable as being Elphidium. 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. Spengler also noted that the septa of foraminifera arced the opposite way from those of nautili and that they lacked a nerve tube.
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. 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.
The protozoan nature of foraminifera was first recognized by Dujardin in 1835. 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.
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".
Geologist Irene Crespin undertook extensive research in this field, publishing some ninety papers—including notable work on foraminifera—as sole author as well as more than twenty in collaboration with other scientists.
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. 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.
Taxonomy
The taxonomic position of the Foraminifera has varied since Schultze in 1854, who referred to as an order, Foraminiferida. Loeblich (1987) and Tappan (1992) reranked Foraminifera as a class as it is now commonly regarded.
The Foraminifera have typically been included in the Protozoa, or in the similar Protoctista or Protist kingdom.eForams-taxonomy Compelling evidence, based primarily on molecular phylogenetics, exists for their belonging to a major group within the Protozoa known as the Rhizaria. 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.
Cavalier-Smith defines the Rhizaria as an infra-kingdom within the kingdom Protozoa.
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. 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.
Taxonomy from Mikhalevich 2013* 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. Some foraminifera lack tests entirely. 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 of the test, and the of the test allow the transfer of material from the pseudopodia to the internal cell and back.