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Foraminifera

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Foraminifera
Temporal range: 542–0 Ma[1] Latest Ediacaran–Recent
Ammonia tepida.jpg
Live Ammonia tepida (Rotaliida)
Scientific classification e
Domain: Eukaryota
Clade: Diaphoretickes
Clade: SAR
Phylum: Retaria
Subphylum: Ectoreta
Infraphylum: Foraminifera
d'Orbigny, 1826
Subdivisions

"Monothalamea"

Tubothalamea

Globothalamea

incertae sedis

Foraminifera (/fəˌræməˈnɪfərə/; Latin for "hole bearers"; informally called "forams") are single-celled organisms, members of a phylum or class of amoeboid protists characterized by streaming granular ectoplasm for catching food and other uses; and commonly an external shell (called a "test") of diverse forms and materials. Tests of chitin (found in some simple genera, and Textularia in particular) are believed to be the most primitive type. Most foraminifera are marine, the majority of which live on or within the seafloor sediment (i.e., are benthic),[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 or multiple chambers, some becoming quite elaborate in structure.[8] These shells are commonly made of calcium carbonate (CaCO
3
) or agglutinated sediment particles. Over 50,000 species are recognized, both living (6,700–10,000)[9][10] and fossil (40,000).[11][12] They are usually less than 1 mm in size, but some are much larger, the largest species reaching up to 20 cm.[13]

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.[14]

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Latin

Latin

Latin is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area around present-day Rome, but through the power of the Roman Republic it became the dominant language in the Italian region and subsequently throughout the Roman Empire. Even after the fall of Western Rome, Latin remained the common language of international communication, science, scholarship and academia in Europe until well into the 18th century, when other regional vernaculars supplanted it in common academic and political usage, and it eventually became a dead language in the modern linguistic definition.

Phylum

Phylum

In biology, a phylum is a level of classification or taxonomic rank below kingdom and above class. Traditionally, in botany the term division has been used instead of phylum, although the International Code of Nomenclature for algae, fungi, and plants accepts the terms as equivalent. Depending on definitions, the animal kingdom Animalia contains about 31 phyla, the plant kingdom Plantae contains about 14 phyla, and the fungus kingdom Fungi contains about 8 phyla. Current research in phylogenetics is uncovering the relationships between phyla, which are contained in larger clades, like Ecdysozoa and Embryophyta.

Class (biology)

Class (biology)

In biological classification, class is a taxonomic rank, as well as a taxonomic unit, a taxon, in that rank. It is a group of related taxonomic orders. Other well-known ranks in descending order of size are life, domain, kingdom, phylum, order, family, genus, and species, with class fitting between phylum and order.

Amoeba

Amoeba

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

Ectoplasm (cell biology)

Ectoplasm (cell biology)

Ectoplasm is the non-granulated outer part of a cell's cytoplasm, while endoplasm is its often granulated inner layer. It is clear, and protects as well as transports things within the cell. Moreover, large numbers of actin filaments frequently occur in the ectoplasm, which form an elastic support for the cell membrane.

Chitin

Chitin

Chitin (C8H13O5N)n ( KY-tin) is a long-chain polymer of N-acetylglucosamine, an amide derivative of glucose. Chitin is probably the second most abundant polysaccharide in nature (behind only cellulose); an estimated 1 billion tons of chitin are produced each year in the biosphere. It is a primary component of cell walls in fungi (especially basidiomycetes and filamentous fungi), the exoskeletons of arthropods such as crustaceans and insects, the radulae, cephalopod beaks and gladii of molluscs and in some nematodes and diatoms. It is also synthesised by at least some fish and lissamphibians. Commercially, chitin is extracted from the shells of crabs, shrimps, shellfish and lobsters, which are major by-products of the seafood industry. The structure of chitin is comparable to cellulose, forming crystalline nanofibrils or whiskers. It is functionally comparable to the protein keratin. Chitin has proved useful for several medicinal, industrial and biotechnological purposes.

Benthos

Benthos

Benthos, also known as benthon, is the community of organisms that live on, in, or near the bottom of a sea, river, lake, or stream, also known as the benthic zone. This community lives in or near marine or freshwater sedimentary environments, from tidal pools along the foreshore, out to the continental shelf, and then down to the abyssal depths.

Plankton

Plankton

Plankton are the diverse collection of organisms found in water that are unable to propel themselves against a current. The individual organisms constituting plankton are called plankters. In the ocean, they provide a crucial source of food to many small and large aquatic organisms, such as bivalves, fish and whales.

Globigerinina

Globigerinina

The Globigerinina is a suborder of foraminiferans that are found as marine plankton. They produce hyaline calcareous tests, and are known as fossils from the Jurassic period onwards. The group has included more than 100 genera and over 400 species, of which about 30 species are extant. One of the most important genera is Globigerina; vast areas of the ocean floor are covered with Globigerina ooze, dominated by the shells of planktonic forms.

Calcium carbonate

Calcium carbonate

Calcium carbonate is a chemical compound with the chemical formula CaCO3. It is a common substance found in rocks as the minerals calcite and aragonite and is the main component of eggshells, gastropod shells, shellfish skeletons and pearls. Things containing much calcium carbonate or resembling it are described as calcareous. Calcium carbonate is the active ingredient in agricultural lime and is created when calcium ions in hard water react with carbonate ions to create limescale. It has medical use as a calcium supplement or as an antacid, but excessive consumption can be hazardous and cause hypercalcemia and digestive issues.

Agglutination (biology)

Agglutination (biology)

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

Fossil

Fossil

A fossil is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in amber, hair, petrified wood and DNA remnants. The totality of fossils is known as the fossil record.

History of study

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]

Earliest known illustration of a foraminifera shell, published by Robert Hooke in his 1665 book Micrographia.
Earliest known illustration of a foraminifera shell, published by Robert Hooke in his 1665 book Micrographia.

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".

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.[21]

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.[22]

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Herodotus

Herodotus

Herodotus was a Greek historian and geographer from the Greek city of Halicarnassus, part of the Persian Empire and a later citizen of Thurii in modern Calabria (Italy). He is known for having written the Histories – a detailed account of the Greco-Persian Wars. Herodotus was the first writer to perform systematic investigation of historical events. He is referred to as "The Father of History", a title conferred on him by the ancient Roman orator Cicero.

Great Pyramid of Giza

Great Pyramid of Giza

The Great Pyramid of Giza is the largest Egyptian pyramid and the tomb of Fourth Dynasty pharaoh Khufu. Built in the early 26th century BC during a period of around 27 years, the pyramid is the oldest of the Seven Wonders of the Ancient World, and the only one to remain largely intact. It is the most famous monument of the Giza pyramid complex, in the Pyramid Fields of the Memphis and its Necropolis UNESCO World Heritage Site, in Giza, Egypt. It is at the most Northern end of the line of the 3 Pyramids of Giza.

Lentil

Lentil

The lentil is an edible legume. It is an annual plant known for its lens-shaped seeds. It is about 40 cm (16 in) tall, and the seeds grow in pods, usually with two seeds in each. As a food crop, the largest producer is Canada, producing 45% of the world’s total lentils.

Micrographia

Micrographia

Micrographia: or Some Physiological Descriptions of Minute Bodies Made by Magnifying Glasses. With Observations and Inquiries Thereupon. is a historically significant book by Robert Hooke about his observations through various lenses. It was the first book to include illustrations of insects and plants as seen through microscopes.

Antonie van Leeuwenhoek

Antonie van Leeuwenhoek

Antonie Philips van Leeuwenhoek was a Dutch microbiologist and microscopist in the Golden Age of Dutch science and technology. A largely self-taught man in science, he is commonly known as "the Father of Microbiology", and one of the first microscopists and microbiologists. Van Leeuwenhoek is best known for his pioneering work in microscopy and for his contributions toward the establishment of microbiology as a scientific discipline.

Elphidium

Elphidium

Elphidium is an abundant genus of foraminifera. Species can be found from coastal regions out to the continental slope, and in all temperature ranges. Like other forams, fossils from different species are used to date rocks. The taxonomy of the species within this genus is disputed due to the high variability of some species.

Cephalopod

Cephalopod

A cephalopod is any member of the molluscan class Cephalopoda such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head, and a set of arms or tentacles modified from the primitive molluscan foot. Fishers sometimes call cephalopods "inkfish", referring to their common ability to squirt ink. The study of cephalopods is a branch of malacology known as teuthology.

Lorenz Spengler

Lorenz Spengler

Lorenz Spengler was a Danish turner and naturalist.

Alcide d'Orbigny

Alcide d'Orbigny

Alcide Charles Victor Marie Dessalines d'Orbigny was a French naturalist who made major contributions in many areas, including zoology, palaeontology, geology, archaeology and anthropology.

Félix Dujardin

Félix Dujardin

Félix Dujardin was a French biologist born in Tours. He is remembered for his research on protozoans and other invertebrates.

Henry Bowman Brady

Henry Bowman Brady

Henry Bowman Brady was a British micropalaeontologist.

Challenger expedition

Challenger expedition

The Challenger expedition of 1872–1876 was a scientific programme that made many discoveries to lay the foundation of oceanography. The expedition was named after the naval vessel that undertook the trip, HMS Challenger.

Taxonomy

"Monothalamids" (paraphyletic)

Lagenida

"Monothalamids"

Tubothalamea

Miliolida

Spirillinida

"Monothalamids"

Xenophyophorea

Globothalamea

"Textulariida" (paraphyletic)

Robertinida

Rotaliida

Phylogeny of Foraminifera following Pawlowski et al. 2013.[22] The monothalamid orders Astrorhizida and Allogromiida are both paraphyletic.

The taxonomic position of the Foraminifera has varied since Schultze in 1854,[23] who referred to as an order, Foraminiferida. Loeblich (1987) and Tappan (1992) reranked Foraminifera as a class[24] as it is now commonly regarded.

The Foraminifera have typically been included in the Protozoa,[25][26][27] or in the similar Protoctista or Protist kingdom.[28][29] Compelling evidence, based primarily on molecular phylogenetics, exists for their belonging to a major group within the Protozoa known as the Rhizaria.[25] 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.[25]

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.[26] 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.[30]

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Lagenida

Lagenida

Lagenida is an order of benthic foraminiferal protists in which the tests (shells) are monolamellar, with walls composed of optically and ultra-structurally radiate calcite, with the crystallographic c-axes perpendicular to the surface. Lagenids first appear in the Upper Silurian and continue to the Recent. They are currently divided into two superfamilies, the older Robuloidacea which range from the Upper Silurian to the Lower Cretaceous (Albian) and the younger Nodosariacea, ranging from the Permian to Recent.

Miliolida

Miliolida

The Miliolida are an order of foraminifera with calcareous, porcelacous tests that are imperforate and commonly have a pseudochitinous lining. Tests are composed of randomly oriented calcite needles that have a high proportion of magnesium along with organic material. Tests lack pores and generally have multiple chambers.

Globothalamea

Globothalamea

Globothalamea comprises a class of multichambered foraminifera based in part on SSU rDNA evidence; the other is Tubothalamea.

Allogromiida

Allogromiida

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

Alfred R. Loeblich Jr

Alfred R. Loeblich Jr

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

Helen Niña Tappan Loeblich

Helen Niña Tappan Loeblich

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

Protozoa

Protozoa

Protozoa are a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Historically, protozoans were regarded as "one-celled animals", because they often possess animal-like behaviours, such as motility and predation, and lack a cell wall, as found in plants and many algae.

Protist

Protist

A protist is any eukaryotic organism that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor, the exclusion of other eukaryotes means that protists do not form a natural group, or clade. Therefore, some protists may be more closely related to animals, plants, or fungi than they are to other protists. However, like the groups algae, invertebrates, and protozoans, the biological category protist is used for convenience. Others classify any unicellular eukaryotic microorganism as a protist. The study of protists is termed protistology.

Kingdom (biology)

Kingdom (biology)

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

Molecular phylogenetics

Molecular phylogenetics

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

Amoeba

Amoeba

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

Phylum

Phylum

In biology, a phylum is a level of classification or taxonomic rank below kingdom and above class. Traditionally, in botany the term division has been used instead of phylum, although the International Code of Nomenclature for algae, fungi, and plants accepts the terms as equivalent. Depending on definitions, the animal kingdom Animalia contains about 31 phyla, the plant kingdom Plantae contains about 14 phyla, and the fungus kingdom Fungi contains about 8 phyla. Current research in phylogenetics is uncovering the relationships between phyla, which are contained in larger clades, like Ecdysozoa and Embryophyta.

Anatomy

Schematic diagram of a live multilocular foraminifera. 1-endoplasm, 2-ectoplasm, 3-chamber, 4-pores, 5-foramen, 6-food vacuole, 7-nucleus, 8-mitochondria, 9-granureticulose pseudopodia, 10-granules, 11- primary aperture, 12-food particle, 13-Golgi apparatus, 14-ribosomes.
Schematic diagram of a live multilocular foraminifera. 1-endoplasm, 2-ectoplasm, 3-chamber, 4-pores, 5-foramen, 6-food vacuole, 7-nucleus, 8-mitochondria, 9-granureticulose pseudopodia, 10-granules, 11- primary aperture, 12-food particle, 13-Golgi apparatus, 14-ribosomes.

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.[24] Some foraminifera lack tests entirely.[32] 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.[33]

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.[34] Foraminifera are unique in having granuloreticulose pseudopodia; that is, 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.[35][33]

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.[24]

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.[36] 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.[37]

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.[37]

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.[37]

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.[38]

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Mollusca

Mollusca

Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks. Around 85,000 extant species of molluscs are recognized. The number of fossil species is estimated between 60,000 and 100,000 additional species. The proportion of undescribed species is very high. Many taxa remain poorly studied.

Coral

Coral

Corals are marine invertebrates within the class Anthozoa of the phylum Cnidaria. They typically form compact colonies of many identical individual polyps. Coral species include the important reef builders that inhabit tropical oceans and secrete calcium carbonate to form a hard skeleton.

Cell membrane

Cell membrane

The cell membrane is a biological membrane that separates and protects the interior of all cells from the outside environment. The cell membrane consists of a lipid bilayer, made up of two layers of phospholipids with cholesterols interspersed between them, maintaining appropriate membrane fluidity at various temperatures. The membrane also contains membrane proteins, including integral proteins that span the membrane and serve as membrane transporters, and peripheral proteins that loosely attach to the outer (peripheral) side of the cell membrane, acting as enzymes to facilitate interaction with the cell's environment. Glycolipids embedded in the outer lipid layer serve a similar purpose. The cell membrane controls the movement of substances in and out of cells and organelles, being selectively permeable to ions and organic molecules. In addition, cell membranes are involved in a variety of cellular processes such as cell adhesion, ion conductivity, and cell signalling and serve as the attachment surface for several extracellular structures, including the cell wall and the carbohydrate layer called the glycocalyx, as well as the intracellular network of protein fibers called the cytoskeleton. In the field of synthetic biology, cell membranes can be artificially reassembled.

Protoplasm

Protoplasm

Protoplasm is the living part of a cell that is surrounded by a plasma membrane. It is a mixture of small molecules such as ions, monosaccharides, amino acid, and macromolecules such as proteins, polysaccharides, lipids, etc.

Pseudopodia

Pseudopodia

A pseudopod or pseudopodium is a temporary arm-like projection of a eukaryotic cell membrane that is emerged in the direction of movement. Filled with cytoplasm, pseudopodia primarily consist of actin filaments and may also contain microtubules and intermediate filaments. Pseudopods are used for motility and ingestion. They are often found in amoebas.

Cytoskeleton

Cytoskeleton

The cytoskeleton is a complex, dynamic network of interlinking protein filaments present in the cytoplasm of all cells, including those of bacteria and archaea. In eukaryotes, it extends from the cell nucleus to the cell membrane and is composed of similar proteins in the various organisms. It is composed of three main components, microfilaments, intermediate filaments and microtubules, and these are all capable of rapid growth or disassembly dependent on the cell's requirements.

Xenophyophorea

Xenophyophorea

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

Radionuclide

Radionuclide

A radionuclide (radioactive nuclide, radioisotope or radioactive isotope) is a nuclide that has excess nuclear energy, making it unstable. This excess energy can be used in one of three ways: emitted from the nucleus as gamma radiation; transferred to one of its electrons to release it as a conversion electron; or used to create and emit a new particle (alpha particle or beta particle) from the nucleus. During those processes, the radionuclide is said to undergo radioactive decay. These emissions are considered ionizing radiation because they are energetic enough to liberate an electron from another atom. The radioactive decay can produce a stable nuclide or will sometimes produce a new unstable radionuclide which may undergo further decay. Radioactive decay is a random process at the level of single atoms: it is impossible to predict when one particular atom will decay. However, for a collection of atoms of a single nuclide the decay rate, and thus the half-life (t1/2) for that collection, can be calculated from their measured decay constants. The range of the half-lives of radioactive atoms has no known limits and spans a time range of over 55 orders of magnitude.

Ecology

Photomicrographs of living planktonic foraminifera [39]   (1) Orbulina universa, (2) Sphaeroidinella dehiscens, (3) Globigerinoides sacculifer, (4) Globigerinoides conglobatus, (5) Globigerinoides ruber (white), (6) Globigerinoides ruber (pink), (7) Globoturborotalita rubescens, (8) Globoturborotalita tenella, (9) Globigerinella calida, (10) Globigerinella siphonifera Type I, (11) Globigerinella siphonifera Type II, (12) Globigerinella adamsi, (13) Globigerina bulloides, (14) Turborotalita quinqueloba, (15) Turborotalita humilis, (16) Hastigerina pelagica, (17) Hastigerinella digitata, (18) Neogloboquadrina incompta, (19) Neogloboquadrina pachyderma, (20) Neogloboquadrina dutertrei, (21) Pulleniatina obliquiloculata, (22) Globorotalia inflata, (23) Globorotalia menardii, (24) Globorotalia scitula, (25) Globorotalia crassaformis, (26) Globorotalia truncatulinoides, (27) Candeina nitida, (28) Globigerinita glutinata, (29) Globigerinita uvula, and (30) Tenuitella fleisheri. Scale bars 200 µm
Photomicrographs of living planktonic foraminifera [39]
(1) Orbulina universa, (2) Sphaeroidinella dehiscens, (3) Globigerinoides sacculifer, (4) Globigerinoides conglobatus, (5) Globigerinoides ruber (white), (6) Globigerinoides ruber (pink), (7) Globoturborotalita rubescens, (8) Globoturborotalita tenella, (9) Globigerinella calida, (10) Globigerinella siphonifera Type I, (11) Globigerinella siphonifera Type II, (12) Globigerinella adamsi, (13) Globigerina bulloides, (14) Turborotalita quinqueloba, (15) Turborotalita humilis, (16) Hastigerina pelagica, (17) Hastigerinella digitata, (18) Neogloboquadrina incompta, (19) Neogloboquadrina pachyderma, (20) Neogloboquadrina dutertrei, (21) Pulleniatina obliquiloculata, (22) Globorotalia inflata, (23) Globorotalia menardii, (24) Globorotalia scitula, (25) Globorotalia crassaformis, (26) Globorotalia truncatulinoides, (27) Candeina nitida, (28) Globigerinita glutinata, (29) Globigerinita uvula, and (30) Tenuitella fleisheri.
Scale bars 200 µm

Modern Foraminifera are primarily marine organisms, but living individuals have been found in brackish, freshwater[34] and even terrestrial habitats.[7] The majority of the species are benthic, and a further 50 morphospecies are planktonic.[35] This count may, however, represent only a fraction of actual diversity, since many genetically distinct species may be morphologically indistinguishable.[40]

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.[24]

The majority of planktonic foraminifera are found in the globigerinina, a lineage within the rotaliida.[22] However, at least one other extant rotaliid lineage, Neogallitellia, seems to have independently evolved a planktonic lifestyle.[41][42] Further, it has been suggested that some Jurassic fossil foraminifera may have also independently evolved a planktonic lifestyle, and may be members of Robertinida.[43]

A number of forams have unicellular algae as endosymbionts, from diverse lineages such as the green algae, red algae, golden algae, diatoms, and dinoflagellates.[35] These mixotrophic foraminifers are particularly common in nutrient-poor oceanic waters.[44] Some forams are kleptoplastic, retaining chloroplasts from ingested algae to conduct photosynthesis.[45]

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.[24] 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.[46] One group, the xenophyophores, has been suggested to farm bacteria within their tests, although studies have failed to find support for this hypothesis.[47] Suspension feeding is also common in the group, and at least some species can take advantage of dissolved organic carbon.[24]

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.[24]

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.[48]

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.[49]

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.[50]

Nonmarine foraminifera have traditionally been neglected in foram research, but recent studies show them to be substantially more diverse than previously known. They are known to inhabit disparate ecological niches, including mosses, rivers, lakes and ponds, wetlands, soils, peat bogs, and sand dunes.[51]

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Benthos

Benthos

Benthos, also known as benthon, is the community of organisms that live on, in, or near the bottom of a sea, river, lake, or stream, also known as the benthic zone. This community lives in or near marine or freshwater sedimentary environments, from tidal pools along the foreshore, out to the continental shelf, and then down to the abyssal depths.

Globigerinina

Globigerinina

The Globigerinina is a suborder of foraminiferans that are found as marine plankton. They produce hyaline calcareous tests, and are known as fossils from the Jurassic period onwards. The group has included more than 100 genera and over 400 species, of which about 30 species are extant. One of the most important genera is Globigerina; vast areas of the ocean floor are covered with Globigerina ooze, dominated by the shells of planktonic forms.

Algae

Algae

Algae is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular microalgae, such as Chlorella, Prototheca and the diatoms, to multicellular forms, such as the giant kelp, a large brown alga which may grow up to 50 metres (160 ft) in length. Most are aquatic and autotrophic and lack many of the distinct cell and tissue types, such as stomata, xylem and phloem that are found in land plants. The largest and most complex marine algae are called seaweeds, while the most complex freshwater forms are the Charophyta, a division of green algae which includes, for example, Spirogyra and stoneworts.

Endosymbiont

Endosymbiont

An endosymbiont or endobiont is any organism that lives within the body or cells of another organism most often, though not always, in a mutualistic relationship. (The term endosymbiosis is from the Greek: ἔνδον endon "within", σύν syn "together" and βίωσις biosis "living".) Examples are nitrogen-fixing bacteria, which live in the root nodules of legumes, single-cell algae inside reef-building corals and bacterial endosymbionts that provide essential nutrients to insects.

Green algae

Green algae

The green algae are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta and Charophyta/Streptophyta. The land plants (Embryophytes) have emerged deep in the Charophyte alga as sister of the Zygnematophyceae. Since the realization that the Embryophytes emerged within the green algae, some authors are starting to include them. The completed clade that includes both green algae and embryophytes is monophyletic and is referred to as the clade Viridiplantae and as the kingdom Plantae. The green algae include unicellular and colonial flagellates, most with two flagella per cell, as well as various colonial, coccoid and filamentous forms, and macroscopic, multicellular seaweeds. There are about 22,000 species of green algae. Many species live most of their lives as single cells, while other species form coenobia (colonies), long filaments, or highly differentiated macroscopic seaweeds.

Golden algae

Golden algae

The Chrysophyceae, usually called chrysophytes, chrysomonads, golden-brown algae or golden algae are a large group of algae, found mostly in freshwater. Golden algae is also commonly used to refer to a single species, Prymnesium parvum, which causes fish kills.

Diatom

Diatom

A diatom is any member of a large group comprising several genera of algae, specifically microalgae, found in the oceans, waterways and soils of the world. Living diatoms make up a significant portion of the Earth's biomass: they generate about 20 to 50 percent of the oxygen produced on the planet each year, take in over 6.7 billion metric tons of silicon each year from the waters in which they live, and constitute nearly half of the organic material found in the oceans. The shells of dead diatoms can reach as much as a half-mile deep on the ocean floor, and the entire Amazon basin is fertilized annually by 27 million tons of diatom shell dust transported by transatlantic winds from the African Sahara, much of it from the Bodélé Depression, which was once made up of a system of fresh-water lakes.

Dinoflagellate

Dinoflagellate

The dinoflagellates are a monophyletic group of single-celled eukaryotes constituting the phylum Dinoflagellata and are usually considered algae. Dinoflagellates are mostly marine plankton, but they also are common in freshwater habitats. Their populations vary with sea surface temperature, salinity, and depth. Many dinoflagellates are photosynthetic, but a large fraction of these are in fact mixotrophic, combining photosynthesis with ingestion of prey.

Chloroplast

Chloroplast

A chloroplast is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it in the energy-storage molecules ATP and NADPH while freeing oxygen from water in the cells. The ATP and NADPH is then used to make organic molecules from carbon dioxide in a process known as the Calvin cycle. Chloroplasts carry out a number of other functions, including fatty acid synthesis, amino acid synthesis, and the immune response in plants. The number of chloroplasts per cell varies from one, in unicellular algae, up to 100 in plants like Arabidopsis and wheat.

Copepod

Copepod

Copepods are a group of small crustaceans found in nearly every freshwater and saltwater habitat. Some species are planktonic, some are benthic, a number of species have parasitic phases, and some continental species may live in limnoterrestrial habitats and other wet terrestrial places, such as swamps, under leaf fall in wet forests, bogs, springs, ephemeral ponds, and puddles, damp moss, or water-filled recesses (phytotelmata) of plants such as bromeliads and pitcher plants. Many live underground in marine and freshwater caves, sinkholes, or stream beds. Copepods are sometimes used as biodiversity indicators.

Cumacea

Cumacea

Cumacea is an order of small marine crustaceans of the superorder Peracarida, occasionally called hooded shrimp or comma shrimp. Their unique appearance and uniform body plan makes them easy to distinguish from other crustaceans. They live in soft-bottoms such as mud and sand, mostly in the marine environment. There are more than 1,500 species of cumaceans formally described. The species diversity of Cumacea increases with depth.

Dissolved organic carbon

Dissolved organic carbon

Dissolved organic carbon (DOC) is the fraction of organic carbon operationally defined as that which can pass through a filter with a pore size typically between 0.22 and 0.7 micrometers. The fraction remaining on the filter is called particulate organic carbon (POC).

Reproduction

The generalized foraminiferal life-cycle involves an alternation between haploid and diploid generations, although they are mostly similar in form.[23][52] 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.[34]

Diagram of a typical foraminiferan life cycle, showing characteristic alternation of generations.
Diagram of a typical foraminiferan life cycle, showing characteristic alternation of generations.

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 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.

Morphs present in the foram life cycle—the megalosphere and the microsphere. The name derives from the size of the proloculus, or first chamber, and as such the microsphere has a larger overall size.
Morphs present in the foram life cycle—the megalosphere and the microsphere. The name derives from the size of the proloculus, or first chamber, and as such the microsphere has a larger overall size.

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.

Fossil nummulitid foraminiferans showing microspheric (larger) and megalospheric individuals (smaller); Eocene of the United Arab Emirates; scale in mm
Fossil nummulitid foraminiferans showing microspheric (larger) and megalospheric individuals (smaller); Eocene of the United Arab Emirates; scale in mm

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.[53][33]

Variations in reproductive mode

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.[33][53][46]

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Cell nucleus

Cell nucleus

The cell nucleus is a membrane-bound organelle found in eukaryotic cells. Eukaryotic cells usually have a single nucleus, but a few cell types, such as mammalian red blood cells, have no nuclei, and a few others including osteoclasts have many. The main structures making up the nucleus are the nuclear envelope, a double membrane that encloses the entire organelle and isolates its contents from the cellular cytoplasm; and the nuclear matrix, a network within the nucleus that adds mechanical support.

Gamete

Gamete

A gamete is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. In species that produce two morphologically distinct types of gametes, and in which each individual produces only one type, a female is any individual that produces the larger type of gamete—called an ovum— and a male produces the smaller type—called a sperm. Sperm cells or spermatozoa are small and motile due to the flagellum, a tail-shaped structure that allows the cell to propel and move. In contrast, each egg cell or ovum is relatively large and non-motile. In short a gamete is an egg cell or a sperm. In animals, ova mature in the ovaries of females and sperm develop in the testes of males. During fertilization, a spermatozoon and ovum unite to form a new diploid organism. Gametes carry half the genetic information of an individual, one ploidy of each type, and are created through meiosis, in which a germ cell undergoes two fissions, resulting in the production of four gametes. In biology, the type of gamete an organism produces determines the classification of its sex.

Flagellum

Flagellum

A flagellum is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates.

Multinucleate

Multinucleate

Multinucleate cells are eukaryotic cells that have more than one nucleus per cell, i.e., multiple nuclei share one common cytoplasm. Mitosis in multinucleate cells can occur either in a coordinated, synchronous manner where all nuclei divide simultaneously or asynchronously where individual nuclei divide independently in time and space. Certain organisms may have a multinuclear stage of their life cycle. For example, slime molds have a vegetative, multinucleate life stage called a plasmodium.

Meiosis

Meiosis

Meiosis is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately result in four cells with only one copy of each chromosome (haploid). Additionally, prior to the division, genetic material from the paternal and maternal copies of each chromosome is crossed over, creating new combinations of code on each chromosome. Later on, during fertilisation, the haploid cells produced by meiosis from a male and female will fuse to create a cell with two copies of each chromosome again, the zygote.

Asexual reproduction

Asexual reproduction

Asexual reproduction is a type of reproduction that does not involve the fusion of gametes or change in the number of chromosomes. The offspring that arise by asexual reproduction from either unicellular or multicellular organisms inherit the full set of genes of their single parent and thus the newly created individual is genetically and physically similar to the parent or an exact clone of the parent. Asexual reproduction is the primary form of reproduction for single-celled organisms such as archaea and bacteria. Many eukaryotic organisms including plants, animals, and fungi can also reproduce asexually. In vertebrates, the most common form of asexual reproduction is parthenogenesis, which is typically used as an alternative to sexual reproduction in times when reproductive opportunities are limited. Komodo dragons and some monitor lizards can also reproduce asexually.

Mitosis

Mitosis

In cell biology, mitosis is a part of the cell cycle in which replicated chromosomes are separated into two new nuclei. Cell division by mitosis gives rise to genetically identical cells in which the total number of chromosomes is maintained. Therefore, mitosis is also known as equational division. In general, mitosis is preceded by S phase of interphase and is often followed by telophase and cytokinesis; which divides the cytoplasm, organelles and cell membrane of one cell into two new cells containing roughly equal shares of these cellular components. The different stages of mitosis altogether define the mitotic (M) phase of an animal cell cycle—the division of the mother cell into two daughter cells genetically identical to each other.

Egg cell

Egg cell

The egg cell, or ovum, is the female reproductive cell, or gamete, in most anisogamous organisms. The term is used when the female gamete is not capable of movement (non-motile). If the male gamete (sperm) is capable of movement, the type of sexual reproduction is also classified as oogamous. A nonmotile female gamete formed in the oogonium of some algae, fungi, oomycetes, or bryophytes is an oosphere. When fertilized the oosphere becomes the oospore.

Cytoplasm

Cytoplasm

In cell biology, the cytoplasm is all of the material within a eukaryotic cell, enclosed by the cell membrane, except for the cell nucleus. The material inside the nucleus and contained within the nuclear membrane is termed the nucleoplasm. The main components of the cytoplasm are cytosol, the organelles, and various cytoplasmic inclusions. The cytoplasm is about 80% water and is usually colorless.

Nummulite

Nummulite

A nummulite is a large lenticular fossil, characterized by its numerous coils, subdivided by septa into chambers. They are the shells of the fossil and present-day marine protozoan Nummulites, a type of foraminiferan. Nummulites commonly vary in diameter from 1.3 cm to 5 cm and are common in Eocene to Miocene marine rocks, particularly around southwest Asia and the Mediterranean. Fossils up to six inches wide are found in the Middle Eocene rocks of Turkey. They are valuable as index fossils.

Eocene

Eocene

The Eocene Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name Eocene comes from the Ancient Greek ἠώς and καινός and refers to the "dawn" of modern ('new') fauna that appeared during the epoch.

Monothalamea

Monothalamea

"Monothalamea" is a grouping of foraminiferans, traditionally consisting of all foraminifera with single-chambered tests. Recent work has shown that the grouping is paraphyletic, and as such does not constitute a natural group; nonetheless, the name "monothalamea" continues to be used by foraminifera workers out of convenience.

Tests

Foraminiferan tests (ventral view)
Foraminiferan tests (ventral view)

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.[54] 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.[33]

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. Tests as fossils are known from as far back as the Ediacaran period,[55] 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.[56] It is estimated that reef Foraminifera generate about 43 million tons of calcium carbonate per year.[57]

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.[58]

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Foraminifera test

Foraminifera test

Foraminiferal tests are the tests of Foraminifera.

Protist shell

Protist shell

Many protists have protective shells or tests, usually made from silica (glass) or calcium carbonate (chalk). Protists are mostly single-celled and microscopic. Their shells are often tough, mineralised forms that resist degradation, and can survive the death of the protist as a microfossil. Although protists are typically very small, they are ubiquitous. Their numbers are such that their shells play a huge part in the formation of ocean sediments and in the global cycling of elements and nutrients.

Test (biology)

Test (biology)

In biology, a test is the hard shell of some spherical marine animals and protists, notably sea urchins and microorganisms such as testate foraminiferans, radiolarians, and testate amoebae. The term is also applied to the covering of scale insects. The related Latin term testa is used for the hard seed coat of plant seeds.

Ediacaran

Ediacaran

The Ediacaran Period is a geological period that spans 96 million years from the end of the Cryogenian Period 635 million years ago (Mya), to the beginning of the Cambrian Period 538.8 Mya. It marks the end of the Proterozoic Eon, and the beginning of the Phanerozoic Eon. It is named after the Ediacara Hills of South Australia.

Nummulite

Nummulite

A nummulite is a large lenticular fossil, characterized by its numerous coils, subdivided by septa into chambers. They are the shells of the fossil and present-day marine protozoan Nummulites, a type of foraminiferan. Nummulites commonly vary in diameter from 1.3 cm to 5 cm and are common in Eocene to Miocene marine rocks, particularly around southwest Asia and the Mediterranean. Fossils up to six inches wide are found in the Middle Eocene rocks of Turkey. They are valuable as index fossils.

Reticulomyxa

Reticulomyxa

Reticulomyxa is a monospecific genus of freshwater foraminiferans. The type species is the unicellular Reticulomyxa filosa. It is found in freshwater environments as well as moist environments, like decomposing matter and damp soils. The heterotrophic naked foraminiferan can feed on microbes as well has larger organisms and is able to be sustained in culture by supplemented nutrients such as wheat germ and oats. The large, multinucleate foraminferan is characteristic for its lack of test and named for the network of connecting pseudopodia surrounding its central body mass. The organism has unique bidirectional cytoplasmic streaming throughout the anastomosing pseudopodia that is some of the fastest reported organelle transport observed. Reticulomyxa was first described in 1949 and is commonly used as a model organism for the unique transport of organelles throughout the cytoplasm of pseudopodia by cytoskeletal mechanisms. Only asexual reproduction of this genus has been observed in culture, but the genome possesses genes related to meiosis suggesting it is capable of sexually reproductive life stages.

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.[59][60]

Due to their non-mineralised tests, "allogromiids" have no fossil record.[59]

The mysterious Paleodictyon has been interpreted as a fossil xenophyophore but this remains controversial.
The mysterious Paleodictyon has been interpreted as a fossil xenophyophore but this remains controversial.

The mysterious vendozoans of the Ediacaran period have been suggested to represent fossil xenophyophores.[61] 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.[62] Other researchers have suggested that the elusive trace fossil Paleodictyon and its relatives may represent a fossil xenophyophore[63] and noted the similarity of the extant xenophyophore Occultammina to the fossil;[64] however, modern examples of Paleodictyon have not been able to clear up the issue and the trace may alternately represent a burrow or a glass sponge.[65] Supporting this notion is the similar habitat of living xenophyophores 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.[64] As of 2017 no definite xenophyophore fossils have been found.[66]

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.[67][55] 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.[68][22] 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.[69]

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.[68]

Cutaway view of a Fusulinid
Cutaway view of a Fusulinid

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 elongated rice-shaped forms.[70][71]

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 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.[33][69][72]

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.[69]

A fossil test from a planktonic globigerininan foraminifera.
A fossil test from a planktonic globigerininan foraminifera.

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.[73][69]

The Robertinida first appear in the fossil record during the Anisian 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.[69]

The first definite Rotaliid fossils do not appear in the fossil record until the Pliensbachian epoch of the Jurassic, following the Triassic-Jurassic event.[74] 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.[69] An additional evolution of planktonic lifestyle occurred in the Miocene or Pliocene, when the rotaliid Neogallitellia independently evolved a planktonic lifestyle.[41][42]

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Neoproterozoic

Neoproterozoic

The Neoproterozoic Era is the unit of geologic time from 1 billion to 538.8 million years ago.

Ediacaran

Ediacaran

The Ediacaran Period is a geological period that spans 96 million years from the end of the Cryogenian Period 635 million years ago (Mya), to the beginning of the Cambrian Period 538.8 Mya. It marks the end of the Proterozoic Eon, and the beginning of the Phanerozoic Eon. It is named after the Ediacara Hills of South Australia.

Allogromiida

Allogromiida

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

Paleodictyon

Paleodictyon

Paleodictyon is a trace fossil, usually interpreted to be a burrow, which appears in the geologic marine record beginning in the Precambrian/Early Cambrian and in modern ocean environments. Paleodictyon were first described by Giuseppe Meneghini in 1850. The origin of the trace fossil is enigmatic and numerous candidates have been proposed.

Diagenesis

Diagenesis

Diagenesis is the process that describes physical and chemical changes in sediments first caused by water-rock interactions, microbial activity, and compaction after their deposition. Increased pressure and temperature only start to play a role as sediments become buried much deeper in the Earth's crust. In the early stages, the transformation of poorly consolidated sediments into sedimentary rock (lithification) is simply accompanied by a reduction in porosity and water expulsion, while their main mineralogical assemblages remain unaltered. As the rock is carried deeper by further deposition above, its organic content is progressively transformed into kerogens and bitumens. The process of diagenesis excludes surface alteration (weathering) and deep metamorphism. There is no sharp boundary between diagenesis and metamorphism, but the latter occurs at higher temperatures and pressures. Hydrothermal solutions, meteoric groundwater, rock porosity, permeability, dissolution/precipitation reactions, and time are all influential factors.

Dickinsonia

Dickinsonia

Dickinsonia is an extinct genus of basal animal that lived during the late Ediacaran period in what is now Australia, China, Russia and Ukraine. The individual Dickinsonia typically resembles a bilaterally symmetrical ribbed oval. Its affinities are presently unknown; its mode of growth is consistent with a stem-group bilaterian affinity, though some have suggested that it belongs to the fungi, or even an "extinct kingdom". The discovery of cholesterol molecules in fossils of Dickinsonia lends support to the idea that Dickinsonia was an animal.

Occultammina

Occultammina

Occultammina is a genus of xenophyophorean foraminifera known from the Atlantic and Pacific oceans. It is notable for being the first known infaunal xenophyophore as well as for being a possible identity for the enigmatic trace fossil Paleodictyon.

Hexactinellid

Hexactinellid

Hexactinellid sponges are sponges with a skeleton made of four- and/or six-pointed siliceous spicules, often referred to as glass sponges. They are usually classified along with other sponges in the phylum Porifera, but some researchers consider them sufficiently distinct to deserve their own phylum, Symplasma. Some experts believe glass sponges are the longest-lived animals on earth; these scientists tentatively estimate a maximum age of up to 15,000 years.

Phanerozoic

Phanerozoic

The Phanerozoic Eon is the current geologic eon in the geologic time scale, and the one during which abundant animal and plant life has existed. It covers 538.8 million years to the present, and it began with the Cambrian Period, when animals first developed hard shells preserved in the fossil record. The time before the Phanerozoic, called the Precambrian, is now divided into the Hadean, Archaean and Proterozoic eons.

Platysolenites

Platysolenites

Platysolenites is a genus of agglutinated foraminifera known from Ediacaran and lower Cambrian assemblages.

Ammodiscus

Ammodiscus

Ammodiscus is a genus in the family Ammodiscidae of textulariid foraminifera. Ammodiscus species are vagile, epibenthic foraminiferans, usually found in seawater with normal salinity.

Ammodiscacea

Ammodiscacea

The Ammodiscacea is a superfamily of foraminifera in the order Textulariida. tests are made of agglutinated grains and consist of a proloculus followed by an enrolled tubular second chamber open at the distal end, that lacks internal septa but which may have growth constrictions.

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.[75] 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.[75] 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[76]

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.[77]

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.[78]

Ten species of planktic foraminifera microfossils from Paleocene-Eocene Thermal Maximum (PETM) sediments in southern Maryland.[79] The scale bars measure 150 microns (0.015 cm). Each specimen is similar in size to a grain of sand.
Ten species of planktic foraminifera microfossils from Paleocene-Eocene Thermal Maximum (PETM) sediments in southern Maryland.[79] The scale bars measure 150 microns (0.015 cm). Each specimen is similar in size to a grain of sand.
Climate change during the last 65 million years as expressed by the oxygen isotope composition of benthic foraminifera. The Paleocene-Eocene thermal maximum is characterized by a brief but prominent excursion, attributed to rapid warming.[80][81][82]
Climate change during the last 65 million years as expressed by the oxygen isotope composition of benthic foraminifera. The Paleocene-Eocene thermal maximum is characterized by a brief but prominent excursion, attributed to rapid warming.[80][81][82]

Since at least 1997, the Paleocene–Eocene thermal maximum (PETM) has been investigated as an analogy for understanding the effects of global warming and of massive carbon inputs to the ocean and atmosphere, including ocean acidification.[83] Humans today emit about 10 Gt of carbon (about 37 Gt CO2e) per year, and at that rate will release a comparable amount to the PETM in about one thousand years. A main difference is that during the PETM the planet was ice-free, as the Drake Passage had not yet opened and the Central American Seaway had not yet closed.[84] Although the PETM is now commonly held to be a case study for global warming and massive carbon emission, the cause, details, and overall significance of the event remain uncertain.[85][86][87][88]

Neoflabellina reticulata from chalk of Rügen, Northeastern Germany. Length:1.2 mm, Age: Upper lower Maastrichtian
Neoflabellina reticulata from chalk of Rügen, Northeastern Germany. Length:1.2 mm, Age: Upper lower Maastrichtian

Foraminifera have significant application in the field of biostratigraphy. Due to their small size and hard shells, foraminifera 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.[89] This application of foraminifera was discovered by Alva C. Ellisor in 1920.[90]

Thin section of a peneroplid foraminiferan from Holocene lagoonal sediment in Rice Bay, San Salvador Island, Bahamas.  Scale bar 100 micrometres
Thin section of a peneroplid foraminiferan from Holocene lagoonal sediment in Rice Bay, San Salvador Island, Bahamas. Scale bar 100 micrometres

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;[91] see δ18O and δ13C. The concentration of trace elements, like strontium (Sr),[92] magnesium (Mg),[93] lithium (Li)[94] and boron (B),[95] 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.

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Ocean Drilling Program

Ocean Drilling Program

The Ocean Drilling Program (ODP) was a multinational effort to explore and study the composition and structure of the Earth's oceanic basins, running from 1985 to 2004. ODP was the successor to the Deep Sea Drilling Project initiated in 1968 by the United States. ODP was an international effort with contributions of Australia, Germany, France, Japan, the United Kingdom and the ESF Consortium for Ocean Drilling (ECOD) including 12 further countries. The program used the drillship JOIDES Resolution on 110 expeditions (legs) to collect about 2,000 deep sea cores from major geological features located in the ocean basins of the world. Drilling discoveries led to further questions and hypotheses, as well as to new disciplines in earth sciences such as the field of paleoceanography.

Jurassic

Jurassic

The Jurassic is a geologic period and stratigraphic system that spanned from the end of the Triassic Period 201.4 million years ago (Mya) to the beginning of the Cretaceous Period, approximately 145 Mya. The Jurassic constitutes the middle period of the Mesozoic Era and is named after the Jura Mountains, where limestone strata from the period were first identified.

Microfossil

Microfossil

A microfossil is a fossil that is generally between 0.001 mm and 1 mm in size, the visual study of which requires the use of light or electron microscopy. A fossil which can be studied with the naked eye or low-powered magnification, such as a hand lens, is referred to as a macrofossil.

Ocean acidification

Ocean acidification

Ocean acidification is the decrease in the pH of the Earth’s ocean. Between 1751 and 2021, the average pH of the ocean surface fell from approximately 8.25 to 8.14. Carbon dioxide emissions from human activities are the primary cause of ocean acidification, with atmospheric carbon dioxide (CO2) levels exceeding 410 ppm (in 2020). CO2 from the atmosphere is absorbed by the oceans. This produces carbonic acid (H2CO3) which dissociates into a bicarbonate ion (HCO−3) and a hydrogen ion (H+). The presence of free hydrogen ions (H+) lowers the pH of the ocean, increasing acidity (this does not mean that seawater is acidic yet; it is still alkaline, with a pH higher than 8).Marine calcifying organisms, such as mollusks, oysters and corals, are especially vulnerable because they rely on calcium carbonate to build shells and skeletons.

Drake Passage

Drake Passage

The Drake Passage is the body of water between South America's Cape Horn, Chile, Argentina and the South Shetland Islands of Antarctica. It connects the southwestern part of the Atlantic Ocean with the southeastern part of the Pacific Ocean and extends into the Southern Ocean. The passage is named after the 16th-century English explorer and privateer Sir Francis Drake.

Central American Seaway

Central American Seaway

The Central American Seaway was a body of water that once separated North America from South America. It formed during the Jurassic during the breakup of the supercontinent Pangaea, and closed when the Isthmus of Panama was formed by volcanic activity in the late Pliocene.

Rügen

Rügen

Rügen is Germany's largest island. It is located off the Pomeranian coast in the Baltic Sea and belongs to the state of Mecklenburg-Western Pomerania.

Maastrichtian

Maastrichtian

The Maastrichtian is, in the ICS geologic timescale, the latest age of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interval from 72.1 to 66 million years ago. The Maastrichtian was preceded by the Campanian and succeeded by the Danian.

Biostratigraphy

Biostratigraphy

Biostratigraphy is the branch of stratigraphy which focuses on correlating and assigning relative ages of rock strata by using the fossil assemblages contained within them. The primary objective of biostratigraphy is correlation, demonstrating that a particular horizon in one geological section represents the same period of time as another horizon at a different section. Fossils within these strata are useful because sediments of the same age can look completely different, due to local variations in the sedimentary environment. For example, one section might have been made up of clays and marls, while another has more chalky limestones. However, if the fossil species recorded are similar, the two sediments are likely to have been laid down around the same time. Ideally these fossils are used to help identify biozones, as they make up the basic biostratigraphy units, and define geological time periods based upon the fossil species found within each section.

Phanerozoic

Phanerozoic

The Phanerozoic Eon is the current geologic eon in the geologic time scale, and the one during which abundant animal and plant life has existed. It covers 538.8 million years to the present, and it began with the Cambrian Period, when animals first developed hard shells preserved in the fossil record. The time before the Phanerozoic, called the Precambrian, is now divided into the Hadean, Archaean and Proterozoic eons.

Radiometric dating

Radiometric dating

Radiometric dating, radioactive dating or radioisotope dating is a technique which is used to date materials such as rocks or carbon, in which trace radioactive impurities were selectively incorporated when they were formed. The method compares the abundance of a naturally occurring radioactive isotope within the material to the abundance of its decay products, which form at a known constant rate of decay. The use of radiometric dating was first published in 1907 by Bertram Boltwood and is now the principal source of information about the absolute age of rocks and other geological features, including the age of fossilized life forms or the age of Earth itself, and can also be used to date a wide range of natural and man-made materials.

Alva C. Ellisor

Alva C. Ellisor

Alva Christine Ellisor (1892–1964) was a geologist and one of the first female stratigraphers in North America.

Modern uses

The oil industry relies heavily on microfossils such as forams to find potential hydrocarbon deposits.[96]

Ammonia beccarii, a benthic foram from the North Sea.
Ammonia beccarii, a benthic foram from the North Sea.

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

Foraminifera have many uses in petroleum exploration and are used routinely to interpret the ages and paleoenvironments of sedimentary strata in oil wells.[97] Agglutinated fossil foraminifera buried deeply in sedimentary basins can be used to estimate thermal maturity, which is a key factor for petroleum generation. The Foraminiferal Colouration Index[98] (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".[99]

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Microfossil

Microfossil

A microfossil is a fossil that is generally between 0.001 mm and 1 mm in size, the visual study of which requires the use of light or electron microscopy. A fossil which can be studied with the naked eye or low-powered magnification, such as a hand lens, is referred to as a macrofossil.

Ammonia (genus)

Ammonia (genus)

Ammonia is a genus of marine foraminifers. It is one of the most abundant foraminifer genera worldwide and occurs in sheltered and shallow marine intertidal environments, sometimes in brackish waters.

North Sea

North Sea

The North Sea lies between Great Britain, Denmark, Norway, Germany, the Netherlands, Belgium and France. An epeiric sea on the European continental shelf, it connects to the Atlantic Ocean through the English Channel in the south and the Norwegian Sea in the north. It is more than 970 kilometres (600 mi) long and 580 kilometres (360 mi) wide, covering 570,000 square kilometres (220,000 sq mi).

Bioindicator

Bioindicator

A bioindicator is any species or group of species whose function, population, or status can reveal the qualitative status of the environment. The most common indicator species are animals. For example, copepods and other small water crustaceans that are present in many water bodies can be monitored for changes that may indicate a problem within their ecosystem. Bioindicators can tell us about the cumulative effects of different pollutants in the ecosystem and about how long a problem may have been present, which physical and chemical testing cannot.

Ocean acidification

Ocean acidification

Ocean acidification is the decrease in the pH of the Earth’s ocean. Between 1751 and 2021, the average pH of the ocean surface fell from approximately 8.25 to 8.14. Carbon dioxide emissions from human activities are the primary cause of ocean acidification, with atmospheric carbon dioxide (CO2) levels exceeding 410 ppm (in 2020). CO2 from the atmosphere is absorbed by the oceans. This produces carbonic acid (H2CO3) which dissociates into a bicarbonate ion (HCO−3) and a hydrogen ion (H+). The presence of free hydrogen ions (H+) lowers the pH of the ocean, increasing acidity (this does not mean that seawater is acidic yet; it is still alkaline, with a pH higher than 8).Marine calcifying organisms, such as mollusks, oysters and corals, are especially vulnerable because they rely on calcium carbonate to build shells and skeletons.

Petroleum

Petroleum

Petroleum, also known as crude oil, or simply oil, is a naturally occurring yellowish-black liquid mixture of mainly hydrocarbons, and is found in geological formations. The name petroleum covers both naturally occurring unprocessed crude oil and petroleum products that consist of refined crude oil. A fossil fuel, petroleum is formed when large quantities of dead organisms, mostly zooplankton and algae, are buried underneath sedimentary rock and subjected to both prolonged heat and pressure.

Foraminiferal Colouration Index

Foraminiferal Colouration Index

The Foraminiferal Colouration Index (FCI) is a tool for assessing the thermal alteration of organic matter buried in sedimentary rock. It uses temperature-controlled colour changes in the organic cement of agglutinated foraminifera (microfossils) to estimate thermal alteration. The method is empirical and based on determination of colour by visual comparison of fossil specimens to the Geological Society of America Rock-Color Chart under a binocular microscope.

Archaeology

Archaeology

Archaeology or archeology is the study of human activity through the recovery and analysis of material culture. The archaeological record consists of artifacts, architecture, biofacts or ecofacts, sites, and cultural landscapes. Archaeology can be considered both a social science and a branch of the humanities. It is usually considered an independent academic discipline, but may also be classified as part of anthropology, history or geography.

Provenance

Provenance

Provenance is the chronology of the ownership, custody or location of a historical object. The term was originally mostly used in relation to works of art but is now used in similar senses in a wide range of fields, including archaeology, paleontology, archives, manuscripts, printed books, the circular economy, and science and computing.

Limestone

Limestone

Limestone is a type of carbonate sedimentary rock which is the main source of the material lime. It is composed mostly of the minerals calcite and aragonite, which are different crystal forms of CaCO3. Limestone forms when these minerals precipitate out of water containing dissolved calcium. This can take place through both biological and nonbiological processes, though biological processes, such as the accumulation of corals and shells in the sea, have likely been more important for the last 540 million years. Limestone often contains fossils which provide scientists with information on ancient environments and on the evolution of life.

Gallery

Source: "Foraminifera", Wikipedia, Wikimedia Foundation, (2023, March 19th), https://en.wikipedia.org/wiki/Foraminifera.

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