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Category Carbonate minerals
(repeating unit)
Strunz classification 5.AB.05
Crystal system Trigonal
Crystal class Hexagonal scalenohedral (3m)
H-M symbol: (3 2/m)
Space group R3c
Unit cell a = 4.9896(2)  Å,
c = 17.0610(11) Å; Z = 6
ColorColorless or white, also gray, yellow, green
Crystal habit Crystalline, granular, stalactitic, concretionary, massive, rhombohedral
Twinning Common by four twin laws
Cleavage Perfect on {1011} three directions with angle of 74° 55' [1]
Fracture Conchoidal
Tenacity Brittle
Mohs scale hardness3 (defining mineral)
Luster Vitreous to pearly on cleavage surfaces
Streak White
Diaphaneity Transparent to translucent
Specific gravity 2.71
Optical propertiesUniaxial (−)
Refractive index nω = 1.640–1.660
nε = 1.486
Birefringence δ = 0.154–0.174
Solubility Soluble in dilute acids
Other characteristicsMay fluoresce red, blue, yellow, and other colors under either SW and LW UV; phosphorescent
References [2] [3] [4]
Crystal structure of calcite Calcite.png
Crystal structure of calcite

Calcite is a carbonate mineral and the most stable polymorph of calcium carbonate (CaCO3). The Mohs scale of mineral hardness, based on scratch hardness comparison, defines value 3 as "calcite".

Carbonate minerals Nickel–Strunz 9 ed mineral class number 5

Carbonate minerals are those minerals containing the carbonate ion, CO32−.

In materials science, polymorphism is the ability of a solid material to exist in more than one form or crystal structure. Polymorphism can potentially be found in any crystalline material including polymers, minerals, and metals, and is related to allotropy, which refers to chemical elements. The complete morphology of a material is described by polymorphism and other variables such as crystal habit, amorphous fraction or crystallographic defects. Polymorphism is relevant to the fields of pharmaceuticals, agrochemicals, pigments, dyestuffs, foods, and explosives.

Calcium carbonate Chemical compound

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


Other polymorphs of calcium carbonate are the minerals aragonite and vaterite. Aragonite will change to calcite over timescales of days or less at temperatures exceeding 300 °C, [5] [6] and vaterite is even less stable.

Aragonite carbonate mineral

Aragonite is a carbonate mineral, one of the three most common naturally occurring crystal forms of calcium carbonate, CaCO3 (the other forms being the minerals calcite and vaterite). It is formed by biological and physical processes, including precipitation from marine and freshwater environments.

Vaterite carbonate mineral

Vaterite is a mineral, a polymorph of calcium carbonate (CaCO3). It was named after the German mineralogist Heinrich Vater. It is also known as mu-calcium carbonate (μ-CaCO3) and has a JCPDS number of 13-192. Vaterite belongs to the hexagonal crystal system, whereas calcite is trigonal and aragonite is orthorhombic.


Calcite is derived from the German Calcit, a term coined in the 19th century from the Latin word for lime, calx (genitive calcis) with the suffix -ite used to name minerals. It is thus etymologically related to chalk. [7]

Lime (material) calcium-containing inorganic mineral

Lime is a calcium-containing inorganic mineral composed primarily of oxides, and hydroxide, usually calcium oxide and/ or calcium hydroxide. It is also the name for calcium oxide which occurs as a product of coal seam fires and in altered limestone xenoliths in volcanic ejecta. The word lime originates with its earliest use as building mortar and has the sense of sticking or adhering.

Chalk A soft, white, porous sedimentary rock made of calcium carbonate

Chalk is a soft, white, porous, sedimentary carbonate rock, a form of limestone composed of the mineral calcite. Calcite is an ionic salt called calcium carbonate or CaCO3. It forms under reasonably deep marine conditions from the gradual accumulation of minute calcite shells (coccoliths) shed from micro-organisms called coccolithophores. Flint (a type of chert) is very common as bands parallel to the bedding or as nodules embedded in chalk. It is probably derived from sponge spicules or other siliceous organisms as water is expelled upwards during compaction. Flint is often deposited around larger fossils such as Echinoidea which may be silicified (i.e. replaced molecule by molecule by flint).

When applied by archaeologists and stone trade professionals, the term alabaster is used not just as in geology and mineralogy, where it is reserved for a variety of gypsum; but also for a similar-looking, translucent variety of fine-grained banded deposit of calcite. [8]

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 and cultural landscapes. Archaeology can be considered both a social science and a branch of the humanities. In North America archaeology is a sub-field of anthropology, while in Europe it is often viewed as either a discipline in its own right or a sub-field of other disciplines.

Alabaster Lightly colored, translucent, and soft calcium minerals, typically gypsum

Alabaster is a mineral or rock that is soft, often used for carving, and is processed for plaster powder. Archaeologists and the stone processing industry use the word differently from geologists. The former use is in a wider sense that includes varieties of two different minerals: the fine-grained massive type of gypsum and the fine-grained banded type of calcite. Geologists define alabaster only as the gypsum type. Chemically, gypsum is a hydrous sulfate of calcium, while calcite is a carbonate of calcium.

Gypsum mineral, calcium sulfate with bounded water

Gypsum is a soft sulfate mineral composed of calcium sulfate dihydrate, with the chemical formula CaSO4·2H2O. It is widely mined and is used as a fertilizer and as the main constituent in many forms of plaster, blackboard chalk and wallboard. A massive fine-grained white or lightly tinted variety of gypsum, called alabaster, has been used for sculpture by many cultures including Ancient Egypt, Mesopotamia, Ancient Rome, the Byzantine Empire and the Nottingham alabasters of Medieval England. Gypsum also crystallizes as beautiful translucent crystals of selenite (mineral). It also forms as an evaporite mineral and as a hydration product of anhydrite.

Unit cell and Miller indices

In publications, two different sets of Miller indices are used to describe directions in calcite crystals - the hexagonal system with three indices h, k, l and the rhombohedral system with four indices h, k, l, i. To add to the complications, there are also two definitions of unit cell for calcite. One, an older "morphological" unit cell, was inferred by measuring angles between faces of crystals and looking for the smallest numbers that fit. Later, a "structural" unit cell was determined using X-ray crystallography. The morphological unit cell has approximate dimensions a = 10 Å and c = 8.5 Å , while for the structural unit cell they are a = 5 Å and c = 17 Å . For the same orientation, c must be multiplied by 4 to convert from morphological to structural units. As an example, the cleavage is given as "perfect on {1 0 1 1}" in morphological coordinates and "perfect on {1 0 1 4}" in structural units. (In hexagonal indices, these are {1 0 1} and {1 0 4}.) Twinning, cleavage and crystal forms are always given in morphological units. [3] [9]

X-ray crystallography Technique used in studying crystal structure

X-ray crystallography (XRC) is a technique used for determining the atomic and molecular structure of a crystal, in which the crystalline structure causes a beam of incident X-rays to diffract into many specific directions. By measuring the angles and intensities of these diffracted beams, a crystallographer can produce a three-dimensional picture of the density of electrons within the crystal. From this electron density, the mean positions of the atoms in the crystal can be determined, as well as their chemical bonds, their crystallographic disorder, and various other information.



Over 800 forms of calcite crystals have been identified. Most common are scalenohedra, with faces in the hexagonal {2 1 1} directions (morphological unit cell) or {2 1 4} directions (structural unit cell); and rhombohedral, with faces in the {1 0 1} or {1 0 4} directions (the most common cleavage plane). [9] Habits include acute to obtuse rhombohedra, tabular forms, prisms, or various scalenohedra. Calcite exhibits several twinning types adding to the variety of observed forms. It may occur as fibrous, granular, lamellar, or compact. A fibrous, efflorescent form is known as lublinite. [10] Cleavage is usually in three directions parallel to the rhombohedron form. Its fracture is conchoidal, but difficult to obtain.

Crystal solid material whose constituent atoms, molecules, or ions are arranged in an ordered pattern extending in all three spatial dimensions

A crystal or crystalline solid is a solid material whose constituents are arranged in a highly ordered microscopic structure, forming a crystal lattice that extends in all directions. In addition, macroscopic single crystals are usually identifiable by their geometrical shape, consisting of flat faces with specific, characteristic orientations. The scientific study of crystals and crystal formation is known as crystallography. The process of crystal formation via mechanisms of crystal growth is called crystallization or solidification.

Prism (geometry) geometric shape, a polyhedron with an n-sided polygonal base

In geometry, a prism is a polyhedron comprising an n-sided polygonal base, a second base which is a translated copy of the first, and n other faces joining corresponding sides of the two bases. All cross-sections parallel to the bases are translations of the bases. Prisms are named for their bases, so a prism with a pentagonal base is called a pentagonal prism. The prisms are a subclass of the prismatoids.

Crystal twinning when two separate crystals share some of the same crystal lattice points in a symmetrical manner

Crystal twinning occurs when two separate crystals share some of the same crystal lattice points in a symmetrical manner. The result is an intergrowth of two separate crystals in a variety of specific configurations. The surface along which the lattice points are shared in twinned crystals is called a composition surface or twin plane.

Scalenohedral faces are chiral and come in pairs with mirror-image symmetry; their growth can be influenced by interaction with chiral biomolecules such as L- and D-amino acids. Rhombohedral faces are achiral. [9]


It has a defining Mohs hardness of 3, a specific gravity of 2.71, and its luster is vitreous in crystallized varieties. Color is white or none, though shades of gray, red, orange, yellow, green, blue, violet, brown, or even black can occur when the mineral is charged with impurities.


Calcite is transparent to opaque and may occasionally show phosphorescence or fluorescence. A transparent variety called Iceland spar is used for optical purposes. Acute scalenohedral crystals are sometimes referred to as "dogtooth spar" while the rhombohedral form is sometimes referred to as "nailhead spar".

Photograph of calcite displaying the characteristic birefringence optical behaviour. Calcite-refraction-property.jpg
Photograph of calcite displaying the characteristic birefringence optical behaviour.

Single calcite crystals display an optical property called birefringence (double refraction). This strong birefringence causes objects viewed through a clear piece of calcite to appear doubled. The birefringent effect (using calcite) was first described by the Danish scientist Rasmus Bartholin in 1669. At a wavelength of ≈590 nm calcite has ordinary and extraordinary refractive indices of 1.658 and 1.486, respectively. [11] Between 190 and 1700 nm, the ordinary refractive index varies roughly between 1.9 and 1.5, while the extraordinary refractive index varies between 1.6 and 1.4. [12]


Calcite, like most carbonates, will dissolve with most forms of acid. Calcite can be either dissolved by groundwater or precipitated by groundwater, depending on several factors including the water temperature, pH, and dissolved ion concentrations. Although calcite is fairly insoluble in cold water, acidity can cause dissolution of calcite and release of carbon dioxide gas. Ambient carbon dioxide, due to its acidity, has a slight solubilizing effect on calcite. Calcite exhibits an unusual characteristic called retrograde solubility in which it becomes less soluble in water as the temperature increases. When conditions are right for precipitation, calcite forms mineral coatings that cement the existing rock grains together or it can fill fractures. When conditions are right for dissolution, the removal of calcite can dramatically increase the porosity and permeability of the rock, and if it continues for a long period of time may result in the formation of caves. On a landscape scale, continued dissolution of calcium carbonate-rich rocks can lead to the expansion and eventual collapse of cave systems, resulting in various forms of karst topography.

Use and applications

One of several calcite or alabaster perfume jars from the tomb of Tutankhamun, d. 1323 BC Alabaster Perfume Jar.jpg
One of several calcite or alabaster perfume jars from the tomb of Tutankhamun, d. 1323 BC

Ancient Egyptians carved many items out of calcite, relating it to their goddess Bast, whose name contributed to the term alabaster because of the close association. Many other cultures have used the material for similar carved objects and applications.

High-grade optical calcite was used in World War II for gun sights, specifically in bomb sights and anti-aircraft weaponry. [13] Also, experiments have been conducted to use calcite for a cloak of invisibility. [14]

Microbiologically precipitated calcite has a wide range of applications, such as soil remediation, soil stabilization and concrete repair.

Calcite, obtained from an 80 kg sample of Carrara marble, [15] is used as the IAEA-603 isotopic standard in mass spectrometry for the calibration of δ18O and δ13C. [16]

Natural occurrence

Calcite is a common constituent of sedimentary rocks, limestone in particular, much of which is formed from the shells of dead marine organisms. Approximately 10% of sedimentary rock is limestone. It is the primary mineral in metamorphic marble. It also occurs in deposits from hot springs as a vein mineral; in caverns as stalactites and stalagmites; and in volcanic or mantle-derived rocks such as carbonatites, kimberlites, or rarely in peridotites.

Calcite is often the primary constituent of the shells of marine organisms, e.g., plankton (such as coccoliths and planktic foraminifera), the hard parts of red algae, some sponges, brachiopods, echinoderms, some serpulids, most bryozoa, and parts of the shells of some bivalves (such as oysters and rudists). Calcite is found in spectacular form in the Snowy River Cave of New Mexico as mentioned above, where microorganisms are credited with natural formations. Trilobites, which became extinct a quarter billion years ago, had unique compound eyes that used clear calcite crystals to form the lenses. [17]

The largest documented single crystal of calcite originated from Iceland, measured 7×7×2 m and 6×6×3 m and weighed about 250 tons. [18]

Formation processes

Calcite formation can proceed by several pathways, from the classical terrace ledge kink model [19] to the crystallization of poorly ordered precursor phases (amorphous calcium carbonate, ACC) via an Ostwald ripening process, or via the agglomeration of nanocrystals. [20]

The crystallization of ACC can occur in two stages: first, the ACC nanoparticles rapidly dehydrate and crystallize to form individual particles of vaterite. Secondly, the vaterite transforms to calcite via a dissolution and reprecipitation mechanism with the reaction rate controlled by the surface area of calcite. [21] The second stage of the reaction is approximately 10 times slower. However, the crystallization of calcite has been observed to be dependent on the starting pH and presence of Mg in solution. [22] A neutral starting pH during mixing promotes the direct transformation of ACC into calcite. Conversely, when ACC forms in a solution that starts with a basic initial pH, the transformation to calcite occurs via metastable vaterite, which forms via a spherulitic growth mechanism. [23] In a second stage this vaterite transforms to calcite via a surface-controlled dissolution and recrystallization mechanism. Mg has a noteworthy effect on both the stability of ACC and its transformation to crystalline CaCO3, resulting in the formation of calcite directly from ACC, as this ion destabilizes the structure of vaterite.

Calcite may form in the subsurface in response to activity of microorganisms, such as during sulfate-dependent anaerobic oxidation of methane, where methane is oxidized and sulfate is reduced by a consortium of methane oxidizers and sulfate reducers, leading to precipitation of calcite and pyrite from the produced bicarbonate and sulfide. These processes can be traced by the specific carbon isotope composition of the calcites, which are extremely depleted in the 13C isotope, by as much as −125 per mil PDB13C). [24]

In Earth history

Calcite seas existed in Earth history when the primary inorganic precipitate of calcium carbonate in marine waters was low-magnesium calcite (lmc), as opposed to the aragonite and high-magnesium calcite (hmc) precipitated today. Calcite seas alternated with aragonite seas over the Phanerozoic, being most prominent in the Ordovician and Jurassic. Lineages evolved to use whichever morph of calcium carbonate was favourable in the ocean at the time they became mineralised, and retained this mineralogy for the remainder of their evolutionary history. [25] Petrographic evidence for these calcite sea conditions consists of calcitic ooids, lmc cements, hardgrounds, and rapid early seafloor aragonite dissolution. [26] The evolution of marine organisms with calcium carbonate shells may have been affected by the calcite and aragonite sea cycle. [27]

Calcite is one of the minerals that has been shown to catalyze an important biological reaction, the formose reaction, and may have had a role in the origin of life. [9] Interaction of its chiral surfaces (see Form) with aspartic acid molecules results in a slight bias in chirality; this is one possible mechanism for the origin of homochirality in living cells. [28]

See also

Related Research Articles

Mineralogy Scientific study of minerals and mineralised artifacts

Mineralogy is a subject of geology specializing in the scientific study of the chemistry, crystal structure, and physical properties of minerals and mineralized artifacts. Specific studies within mineralogy include the processes of mineral origin and formation, classification of minerals, their geographical distribution, as well as their utilization.

Strontianite carbonate mineral

Strontianite (SrCO3) is an important raw material for the extraction of strontium. It is a rare carbonate mineral and one of only a few strontium minerals. It is a member of the aragonite group.

Dolomite carbonate mineral - CaMg(CO₃)₂

Dolomite is an anhydrous carbonate mineral composed of calcium magnesium carbonate, ideally CaMg(CO3)2. The term is also used for a sedimentary carbonate rock composed mostly of the mineral dolomite. An alternative name sometimes used for the dolomitic rock type is dolostone.

Rhodochrosite manganese carbonate mineral

Rhodochrosite is a manganese carbonate mineral with chemical composition MnCO3. In its (rare) pure form, it is typically a rose-red color, but impure specimens can be shades of pink to pale brown. It streaks white, and its Mohs hardness varies between 3.5 and 4. Its specific gravity is between 3.5 and 3.7. It crystallizes in the trigonal system, and cleaves with rhombohedral carbonate cleavage in three directions. Crystal twinning often is present. It is transparent to translucent with refractive indices of =1.814 to 1.816, =1.596 to 1.598. It is often confused with the manganese silicate, rhodonite, but is distinctly softer.

Speleothem A structure formed in a cave by the deposition of minerals from water

Speleothems, commonly known as cave formations, are secondary mineral deposits formed in a cave. Speleothems typically form in limestone or dolostone solutional caves. The term "speleothem" as first introduced by Moore (1952), is derived from the Greek words spēlaion "cave" + théma "deposit". The definition of "speleothem" in most publications, specifically excludes secondary mineral deposits in mines, tunnels and on man-made structures. Hill and Forti more concisely defined "secondary minerals" which create speleothems in caves as;

A "secondary" mineral is one which is derived by a physicochemical reaction from a primary mineral in bedrock or detritus, and/or deposited because of a unique set of conditions in a cave; i.e., the cave environment has influenced the mineral's deposition.

Ooid sedimentary grains

Ooids are small, spheroidal, "coated" (layered) sedimentary grains, usually composed of calcium carbonate, but sometimes made up of iron- or phosphate-based minerals. Ooids usually form on the sea floor, most commonly in shallow tropical seas. After being buried under additional sediment, these ooid grains can be cemented together to form a sedimentary rock called an oolite. Oolites usually consist of calcium carbonate; these belong to the limestone rock family. Pisoids are similar to ooids, but are larger than 2 mm in diameter, often considerably larger, as with the pisoids in the hot springs at Carlsbad in the Czech Republic.

Alstonite carbonate mineral

Alstonite, also known as bromlite, is a low temperature hydrothermal mineral that is a rare double carbonate of calcium and barium with the formula BaCa(CO
, sometimes with some strontium. Barytocalcite and paralstonite have the same formula but different structures, so these three minerals are said to be trimorphous. Alstonite is triclinic but barytocalcite is monoclinic and paralstonite is trigonal. The species was named bromlite by Thomas Thomson in 1837 after the Bromley-Hill mine, and alstonite by August Breithaupt of the Freiberg Mining Academy in 1841, after Alston, Cumbria, the base of operations of the mineral dealer from whom the first samples were obtained by Thomson in 1834. Both of these names have been in common use.

Barytocalcite carbonate mineral

Barytocalcite is an anhydrous barium calcium carbonate mineral with the chemical formula BaCa(CO3)2. It is trimorphous with alstonite and paralstonite, that is to say the three minerals have the same formula but different structures. Baryte and quartz pseudomorphs after barytocalcite have been observed.

Leadhillite carbonate-sulfate mineral

Leadhillite is a lead sulfate carbonate hydroxide mineral, often associated with anglesite. It has the formula Pb4SO4(CO3)2(OH)2. Leadhillite crystallises in the monoclinic system, but develops pseudo-hexagonal forms due to crystal twinning. It forms transparent to translucent variably coloured crystals with an adamantine lustre. It is quite soft with a Mohs hardness of 2.5 and a relatively high specific gravity of 6.26 to 6.55.


Anthodites (Greek ἄνθος ánthos, “flower”, -ode, adjectival combining form, -ite adjectival suffix) are speleothems (cave formations) composed of long needle-like crystals situated in clusters which radiate outward from a common base. The "needles" may be quill-like or feathery. Most anthodites are made of the mineral aragonite (a variety of calcium carbonate, CaCO3), although some are composed of gypsum (CaSO4·2H2O).

Kutnohorite carbonate mineral

Kutnohorite is a rare calcium manganese carbonate mineral with magnesium and iron that is a member of the dolomite group. It forms a series with dolomite, and with ankerite. The end member formula is CaMn2+(CO3)2, but Mg2+ and Fe2+ commonly substitute for Mn2+, with the Mn content varying from 38% to 84%, so the formula Ca(Mn2+,Mg,Fe2+)(CO3)2 better represents the species. It was named by Professor Bukowsky in 1901 after the type locality of Kutná Hora, Bohemia, in the Czech Republic. It was originally spelt “kutnahorite” but “kutnohorite” is the current IMA-approved spelling.

Ikaite carbonate mineral

Ikaite is the mineral name for the hexahydrate of calcium carbonate, CaCO3·6H2O. Ikaite tends to form very steep or spiky pyramidal crystals, often radially arranged, of varied sizes from thumbnail size aggregates to gigantic salient spurs. It is only found in a metastable state, and decomposes rapidly by losing most of its water content once removed from near-freezing water. This 'melting mineral' is more commonly known through its pseudomorphs.

Monohydrocalcite carbonate mineral

Monohydrocalcite is a mineral that is a hydrous form of calcium carbonate, CaCO3·H2O. It was formerly also known by the name hydrocalcite, which is now discredited by the IMA. It is a trigonal mineral which is white when pure. Monohydrocalcite is not a common rock-forming mineral, but is frequently associated with other calcium and magnesium carbonate minerals, such as calcite, aragonite, lansfordite, and nesquehonite.

Calcite sea Sea chemistry favouring low-magnesium calcite as the inorganic calcium carbonate precipitate

A calcite sea is one in which low-magnesium calcite is the primary inorganic marine calcium carbonate precipitate. An aragonite sea is the alternate seawater chemistry in which aragonite and high-magnesium calcite are the primary inorganic carbonate precipitates. The Early Paleozoic and the Middle to Late Mesozoic oceans were predominantly calcite seas, whereas the Middle Paleozoic through the Early Mesozoic and the Cenozoic are characterized by aragonite seas.

Aragonite sea Chemical conditions of the sea favouring aragonite deposition

An aragonite sea contains aragonite and high-magnesium calcite as the primary inorganic calcium carbonate precipitates. The chemical conditions of the seawater must be notably high in magnesium content relative to calcium for an aragonite sea to form. This is in contrast to a calcite sea in which seawater low in magnesium content relative to calcium favors the formation of low-magnesium calcite as the primary inorganic marine calcium carbonate precipitate.

Amorphous calcium carbonate

Amorphous calcium carbonate (ACC) is the amorphous and least stable polymorph of calcium carbonate. ACC is extremely unstable under normal conditions and is found naturally in taxa as wide-ranging as sea urchins, corals, mollusks, and foraminifera. It is usually found as a monohydrate, holding the chemical formula CaCO3·H2O; however, it can also exist in a dehydrated state, CaCO3. ACC has been known to science for over 100 years when a non-diffraction pattern of calcium carbonate was discovered by Sturcke Herman, exhibiting its poorly-ordered nature.

Talmessite arsenate mineral

Talmessite is a hydrated calcium magnesium arsenate, often with significant amounts of cobalt or nickel. It was named in 1960 for the type locality, the Talmessi mine, Anarak district, Iran. It forms a series with β-Roselite, where cobalt replaces some of the magnesium, and with gaitite, where zinc replaces the magnesium. All these minerals are members of the fairfieldite group. Talmessite is dimorphic with wendwilsonite.

Marine biogenic calcification is the process by which marine organisms such as oysters and clams form calcium carbonate. Seawater is full of dissolved compounds, ions and nutrients that organisms can utilize for energy and, in the case of calcification, to build shells and outer structures. Calcifying organisms in the ocean include molluscs, foraminifera, coccolithophores, crustaceans, echinoderms such as sea urchins, and corals. The shells and skeletons produced from calcification have important functions for the physiology and ecology of the organisms that create them.


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Further reading