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Exoskeleton

From Wikipedia, in a visual modern way
Part of a series related to
Biomineralization
General
Exoskeletons (shells)
Endoskeletons (bones)
Teeth, scales, tusks etc
Calcification
Silicification
Other forms
Related
"Robot exoskeleton" redirects here. For the type of machine, see powered exoskeleton.
The discarded exoskeleton (exuviae) of dragonfly nymph
The discarded exoskeleton (exuviae) of dragonfly nymph
Exoskeleton of cicada attached to a Tridax procumbens (colloquially known as the tridax daisy)
Exoskeleton of cicada attached to a Tridax procumbens (colloquially known as the tridax daisy)

An exoskeleton (from Greek έξω éxō "outer"[1] and σκελετός skeletós "skeleton"[2][3]) is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton (endoskeleton) in for example, a human. In usage, some of the larger kinds of exoskeletons are known as "shells". Examples of exoskeletons within animals include the arthropod exoskeleton shared by chelicerates, myriapods, crustaceans, and insects, as well as the shell of certain sponges and the mollusc shell shared by snails, clams, tusk shells, chitons and nautilus. Some animals, such as the turtle, have both an endoskeleton and an exoskeleton.

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Skeleton

Skeleton

A skeleton is the structural frame that supports the body of most animals. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside the body, and the hydroskeleton, a flexible internal skeleton supported by fluid pressure. Vertebrates are animals with a vertebral column, and their skeletons are typically composed of bone and cartilage. Invertebrates are animals that lack a vertebral column. The skeletons of invertebrates vary, including hard exoskeleton shells, plated endoskeletons, or spicules. Cartilage is a rigid connective tissue that is found in the skeletal systems of vertebrates and invertebrates.

Endoskeleton

Endoskeleton

An endoskeleton is an internal support structure of an animal, composed of mineralized tissue.

Human skeleton

Human skeleton

The human skeleton is the internal framework of the human body. It is composed of around 270 bones at birth – this total decreases to around 206 bones by adulthood after some bones get fused together. The bone mass in the skeleton makes up about 14% of the total body weight and reaches maximum mass between the ages of 25 and 30. The human skeleton can be divided into the axial skeleton and the appendicular skeleton. The axial skeleton is formed by the vertebral column, the rib cage, the skull and other associated bones. The appendicular skeleton, which is attached to the axial skeleton, is formed by the shoulder girdle, the pelvic girdle and the bones of the upper and lower limbs.

Armour (anatomy)

Armour (anatomy)

Armour or armor in animals is external or superficial protection against attack by predators, formed as part of the body, usually through the hardening of body tissues, outgrowths or secretions. It has therefore mostly developed in 'prey' species.

Arthropod exoskeleton

Arthropod exoskeleton

Arthropods are covered with a tough, resilient integument or exoskeleton of chitin. Generally the exoskeleton will have thickened areas in which the chitin is reinforced or stiffened by materials such as minerals or hardened proteins. This happens in parts of the body where there is a need for rigidity or elasticity. Typically the mineral crystals, mainly calcium carbonate, are deposited among the chitin and protein molecules in a process called biomineralization. The crystals and fibres interpenetrate and reinforce each other, the minerals supplying the hardness and resistance to compression, while the chitin supplies the tensile strength. Biomineralization occurs mainly in crustaceans. In insects and arachnids, the main reinforcing materials are various proteins hardened by linking the fibres in processes called sclerotisation and the hardened proteins are called sclerotin. The dorsal tergum, ventral sternum, and the lateral pleura form the hardened plates or sclerites of a typical body segment.

Crustacean

Crustacean

Crustaceans form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group can be treated as a subphylum under the clade Mandibulata. It is now well accepted that the hexapods emerged deep in the Crustacean group, with the completed group referred to as Pancrustacea. Some crustaceans are more closely related to insects and the other hexapods than they are to certain other crustaceans.

Insect

Insect

Insects are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body, three pairs of jointed legs, compound eyes and one pair of antennae. Their blood is not totally contained in vessels; some circulates in an open cavity known as the haemocoel. Insects are the most diverse group of animals; they include more than a million described species and represent more than half of all known living organisms. The total number of extant species is estimated at between six and ten million; potentially over 90% of the animal life forms on Earth are insects. Insects may be found in nearly all environments, although only a small number of species reside in the oceans, which are dominated by another arthropod group, crustaceans, which recent research has indicated insects are nested within.

Mollusc shell

Mollusc shell

The mollusc shell is typically a calcareous exoskeleton which encloses, supports and protects the soft parts of an animal in the phylum Mollusca, which includes snails, clams, tusk shells, and several other classes. Not all shelled molluscs live in the sea; many live on the land and in freshwater.

Snail

Snail

A snail is a shelled gastropod. The name is most often applied to land snails, terrestrial pulmonate gastropod molluscs. However, the common name snail is also used for most of the members of the molluscan class Gastropoda that have a coiled shell that is large enough for the animal to retract completely into. When the word "snail" is used in this most general sense, it includes not just land snails but also numerous species of sea snails and freshwater snails. Gastropods that naturally lack a shell, or have only an internal shell, are mostly called slugs, and land snails that have only a very small shell are often called semi-slugs.

Bivalvia

Bivalvia

Bivalvia, in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, bivalves have no head and they lack some usual molluscan organs, like the radula and the odontophore. The class includes the clams, oysters, cockles, mussels, scallops, and numerous other families that live in saltwater, as well as a number of families that live in freshwater. The majority are filter feeders. The gills have evolved into ctenidia, specialised organs for feeding and breathing. Most bivalves bury themselves in sediment, where they are relatively safe from predation. Others lie on the sea floor or attach themselves to rocks or other hard surfaces. Some bivalves, such as the scallops and file shells, can swim. The shipworms bore into wood, clay, or stone and live inside these substances.

Chiton

Chiton

Chitons are marine molluscs of varying size in the class Polyplacophora, formerly known as Amphineura. About 940 extant and 430 fossil species are recognized.

Nautilus

Nautilus

The nautilus is a pelagic marine mollusc of the cephalopod family Nautilidae. The nautilus is the sole extant family of the superfamily Nautilaceae and of its smaller but near equal suborder, Nautilina.

Role

Exoskeletons contain rigid and resistant components that fulfill a set of functional roles in many animals including protection, excretion, sensing, support, feeding and acting as a barrier against desiccation in terrestrial organisms. Exoskeletons have a role in defense from pests and predators, support and in providing an attachment framework for musculature.[4]

Arthropod exoskeletons contain chitin; the addition of calcium carbonate makes them harder and stronger, at the price of increased weight.[5] Ingrowths of the arthropod exoskeleton known as apodemes serve as attachment sites for muscles. These structures are composed of chitin and are approximately six times stronger and twice the stiffness of vertebrate tendons. Similar to tendons, apodemes can stretch to store elastic energy for jumping, notably in locusts.[6] Calcium carbonates constitute the shells of molluscs, brachiopods, and some tube-building polychaete worms. Silica forms the exoskeleton in the microscopic diatoms and radiolaria. One species of mollusc, the scaly-foot gastropod, even makes use of the iron sulfides greigite and pyrite.

Some organisms, such as some foraminifera, agglutinate exoskeletons by sticking grains of sand and shell to their exterior. Contrary to a common misconception, echinoderms do not possess an exoskeleton, as their test is always contained within a layer of living tissue.

Exoskeletons have evolved independently many times; 18 lineages evolved calcified exoskeletons alone.[7] Further, other lineages have produced tough outer coatings analogous to an exoskeleton, such as some mammals. This coating is constructed from bone in the armadillo, and hair in the pangolin. The armor of reptiles like turtles and dinosaurs like Ankylosaurs is constructed of bone; crocodiles have bony scutes and horny scales.

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Desiccation

Desiccation

Desiccation is the state of extreme dryness, or the process of extreme drying. A desiccant is a hygroscopic substance that induces or sustains such a state in its local vicinity in a moderately sealed container.

Arthropod exoskeleton

Arthropod exoskeleton

Arthropods are covered with a tough, resilient integument or exoskeleton of chitin. Generally the exoskeleton will have thickened areas in which the chitin is reinforced or stiffened by materials such as minerals or hardened proteins. This happens in parts of the body where there is a need for rigidity or elasticity. Typically the mineral crystals, mainly calcium carbonate, are deposited among the chitin and protein molecules in a process called biomineralization. The crystals and fibres interpenetrate and reinforce each other, the minerals supplying the hardness and resistance to compression, while the chitin supplies the tensile strength. Biomineralization occurs mainly in crustaceans. In insects and arachnids, the main reinforcing materials are various proteins hardened by linking the fibres in processes called sclerotisation and the hardened proteins are called sclerotin. The dorsal tergum, ventral sternum, and the lateral pleura form the hardened plates or sclerites of a typical body segment.

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.

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.

Elastic energy

Elastic energy

Elastic energy is the mechanical potential energy stored in the configuration of a material or physical system as it is subjected to elastic deformation by work performed upon it. Elastic energy occurs when objects are impermanently compressed, stretched or generally deformed in any manner. Elasticity theory primarily develops formalisms for the mechanics of solid bodies and materials. The elastic potential energy equation is used in calculations of positions of mechanical equilibrium. The energy is potential as it will be converted into other forms of energy, such as kinetic energy and sound energy, when the object is allowed to return to its original shape (reformation) by its elasticity.

Locust

Locust

Locusts are various species of short-horned grasshoppers in the family Acrididae that have a swarming phase. These insects are usually solitary, but under certain circumstances they become more abundant and change their behaviour and habits, becoming gregarious. No taxonomic distinction is made between locust and grasshopper species; the basis for the definition is whether a species forms swarms under intermittently suitable conditions; this has evolved independently in multiple lineages, comprising at least 18 genera in 5 different acridid subfamilies.

Brachiopod

Brachiopod

Brachiopods, phylum Brachiopoda, are a phylum of trochozoan animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, while the front can be opened for feeding or closed for protection. Two major categories are traditionally recognized, articulate and inarticulate brachiopods. The word "articulate" is used to describe the tooth-and-groove structures of the valve-hinge which is present in the articulate group, and absent from the inarticulate group. This is the leading diagnostic skeletal feature, by which the two main groups can be readily distinguished as fossils. Articulate brachiopods have toothed hinges and simple, vertically-oriented opening and closing muscles. Conversely, inarticulate brachiopods have weak, untoothed hinges and a more complex system of vertical and oblique (diagonal) muscles used to keep the two valves aligned. In many brachiopods, a stalk-like pedicle projects from an opening near the hinge of one of the valves, known as the pedicle or ventral valve. The pedicle, when present, keeps the animal anchored to the seabed but clear of sediment which would obstruct the opening.

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.

Greigite

Greigite

Greigite is an iron sulfide mineral with the chemical formula Fe2+Fe3+2S4. It is the sulfur equivalent of the iron oxide magnetite (Fe3O4). It was first described in 1964 for an occurrence in San Bernardino County, California, and named after the mineralogist and physical chemist Joseph W. Greig (1895–1977).

Foraminifera

Foraminifera

Foraminifera 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 of diverse forms and materials. Tests of chitin are believed to be the most primitive type. Most foraminifera are marine, the majority of which live on or within the seafloor sediment, while a smaller number float in the water column at various depths, which belong to the suborder Globigerinina. Fewer are known from freshwater or brackish conditions, and some very few (nonaquatic) soil species have been identified through molecular analysis of small subunit ribosomal DNA.

Agglutination (biology)

Agglutination (biology)

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

Echinoderm

Echinoderm

An echinoderm is any member of the phylum Echinodermata. The adults are recognisable by their radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the sea lilies or "stone lilies". Adult echinoderms are found on the sea bed at every ocean depth, from the intertidal zone to the abyssal zone. The phylum contains about 7,000 living species, making it the second-largest grouping of deuterostomes, after the chordates. Echinoderms are the largest entirely marine phylum. The first definitive echinoderms appeared near the start of the Cambrian.

Growth

Main article: Ecdysis

Since exoskeletons are rigid, they present some limits to growth. Organisms with open shells can grow by adding new material to the aperture of their shell, as is the case in snails, bivalves and other molluscans. A true exoskeleton, like that found in arthropods, must be shed (moulted) when it is outgrown.[8] A new exoskeleton is produced beneath the old one. As the old one is shed, the new skeleton is soft and pliable. The animal will typically stay in a den or burrow for this time, as it is quite vulnerable during this period. Once at least partially set, the organism will plump itself up to try to expand the exoskeleton. The new exoskeleton is still capable of growing to some degree, however. Animals of the order arthropoda, like lizards, amphibians, and many other animals that shed their skin, are indeterminate growers.[9] Animals that are indeterminate growers grow in size continually throughout their life because, in this case, their exoskeleton is always being replaced. Failure to shed the exoskeleton once outgrown can result in the animal being suffocated within its own shell, and will stop subadults from reaching maturity, thus preventing them from reproducing. This is the mechanism behind some insect pesticides, such as Azadirachtin.[10]

Paleontological significance

Borings in exoskeletons can provide evidence of animal behavior. In this case, boring sponges attacked this hard clam shell after the death of the clam, producing the trace fossil Entobia.
Borings in exoskeletons can provide evidence of animal behavior. In this case, boring sponges attacked this hard clam shell after the death of the clam, producing the trace fossil Entobia.

Exoskeletons, as hard parts of organisms, are greatly useful in assisting preservation of organisms, whose soft parts usually rot before they can be fossilized. Mineralized exoskeletons can be preserved "as is", as shell fragments, for example. The possession of an exoskeleton permits a couple of other routes to fossilization. For instance, the tough layer can resist compaction, allowing a mold of the organism to be formed underneath the skeleton, which may later decay.[11] Alternatively, exceptional preservation may result in chitin being mineralized, as in the Burgess Shale,[12] or transformed to the resistant polymer keratin, which can resist decay and be recovered.

However, our dependence on fossilized skeletons also significantly limits our understanding of evolution. Only the parts of organisms that were already mineralized are usually preserved, such as the shells of molluscs. It helps that exoskeletons often contain "muscle scars", marks where muscles have been attached to the exoskeleton, which may allow the reconstruction of much of an organism's internal parts from its exoskeleton alone.[11] The most significant limitation is that, although there are 30-plus phyla of living animals, two-thirds of these phyla have never been found as fossils, because most animal species are soft-bodied and decay before they can become fossilized.[13]

Mineralized skeletons first appear in the fossil record shortly before the base of the Cambrian period, 550 million years ago. The evolution of a mineralized exoskeleton is seen by some as a possible driving force of the Cambrian explosion of animal life, resulting in a diversification of predatory and defensive tactics. However, some Precambrian (Ediacaran) organisms produced tough outer shells[11] while others, such as Cloudina, had a calcified exoskeleton.[14] Some Cloudina shells even show evidence of predation, in the form of borings.[14]

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Sponge

Sponge

Sponges, the members of the phylum Porifera, are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate through them, consisting of jelly-like mesohyl sandwiched between two thin layers of cells.

Hard clam

Hard clam

The hard clam, also known as the round clam, hard-shell clam, or the quahog, is an edible marine bivalve mollusk that is native to the eastern shores of North America and Central America from Prince Edward Island to the Yucatán Peninsula. It is one of many unrelated edible bivalves that in the United States are frequently referred to simply as clams. Older literature sources may use the systematic name Venus mercenaria; this species is in the family Veneridae, the venus clams.

Entobia

Entobia

Entobia is a trace fossil in a hard substrate formed by sponges as a branching network of galleries, often with regular enlargements termed chambers. Apertural canals connect the outer surface of the substrate to the chambers and galleries so the sponge can channel water through its tissues for filter feeding. The fossil ranges from the Devonian to the Recent.

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.

Lagerstätte

Lagerstätte

A Lagerstätte is a sedimentary deposit that exhibits extraordinary fossils with exceptional preservation—sometimes including preserved soft tissues. These formations may have resulted from carcass burial in an anoxic environment with minimal bacteria, thus delaying the decomposition of both gross and fine biological features until long after a durable impression was created in the surrounding matrix. Lagerstätten span geological time from the Neoproterozoic era to the present. Worldwide, some of the best examples of near-perfect fossilization are the Cambrian Maotianshan shales and Burgess Shale, the Silurian Waukesha Biota, the Devonian Hunsrück Slates and Gogo Formation, the Carboniferous Mazon Creek, the Jurassic Posidonia Shale and Solnhofen Limestone, the Cretaceous Yixian, Santana, and Agua Nueva formations, the Eocene Green River Formation, the Miocene Foulden Maar and Ashfall Fossil Beds, the Pliocene Gray Fossil Site, the Pleistocene Naracoorte Caves, the La Brea Tar Pits, and the Tanis Fossil Site.

Burgess Shale

Burgess Shale

The Burgess Shale is a fossil-bearing deposit exposed in the Canadian Rockies of British Columbia, Canada. It is famous for the exceptional preservation of the soft parts of its fossils. At 508 million years old, it is one of the earliest fossil beds containing soft-part imprints.

Keratin

Keratin

Keratin is one of a family of structural fibrous proteins also known as scleroproteins. Alpha-keratin (α-keratin) is a type of keratin found in vertebrates. It is the key structural material making up scales, hair, nails, feathers, horns, claws, hooves, and the outer layer of skin among vertebrates. Keratin also protects epithelial cells from damage or stress. Keratin is extremely insoluble in water and organic solvents. Keratin monomers assemble into bundles to form intermediate filaments, which are tough and form strong unmineralized epidermal appendages found in reptiles, birds, amphibians, and mammals. Excessive keratinization participate in fortification of certain tissues such as in horns of cattle and rhinos, and armadillos' osteoderm. The only other biological matter known to approximate the toughness of keratinized tissue is chitin. Keratin comes in two types, the primitive, softer forms found in all vertebrates and harder, derived forms found only among sauropsids.

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.

Cambrian explosion

Cambrian explosion

The Cambrian explosion, Cambrian radiation, Cambrian diversification, or the Biological Big Bang refers to an interval of time approximately 538.8 million years ago in the Cambrian Period when practically all major animal phyla started appearing in the fossil record. It lasted for about 13 – 25 million years and resulted in the divergence of most modern metazoan phyla. The event was accompanied by major diversification in other groups of organisms as well.

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.

Evolution

Further information: Small shelly fauna

On the whole, the fossil record only contains mineralised exoskeletons, since these are by far the most durable. Since most lineages with exoskeletons are thought to have started out with a non-mineralised exoskeleton which they later mineralised, this makes it difficult to comment on the very early evolution of each lineage's exoskeleton. It is known, however, that in a very short course of time, just before the Cambrian period, exoskeletons made of various materials – silica, calcium phosphate, calcite, aragonite, and even glued-together mineral flakes – sprang up in a range of different environments.[15] Most lineages adopted the form of calcium carbonate which was stable in the ocean at the time they first mineralised, and did not change from this mineral morph - even when it became the less favorable.[7]

Some Precambrian (Ediacaran) organisms produced tough but non-mineralized outer shells,[11] while others, such as Cloudina, had a calcified exoskeleton,[14] but mineralized skeletons did not become common until the beginning of the Cambrian period, with the rise of the "small shelly fauna". Just after the base of the Cambrian, these miniature fossils become diverse and abundant – this abruptness may be an illusion, since the chemical conditions which preserved the small shellies appeared at the same time.[16] Most other shell-forming organisms appear during the Cambrian period, with the Bryozoans being the only calcifying phylum to appear later, in the Ordovician. The sudden appearance of shells has been linked to a change in ocean chemistry which made the calcium compounds of which the shells are constructed stable enough to be precipitated into a shell. However this is unlikely to be a sufficient cause, as the main construction cost of shells is in creating the proteins and polysaccharides required for the shell's composite structure, not in the precipitation of the mineral components.[4] Skeletonization also appeared at almost exactly the same time that animals started burrowing to avoid predation, and one of the earliest exoskeletons was made of glued-together mineral flakes, suggesting that skeletonization was likewise a response to increased pressure from predators.[15]

Ocean chemistry may also control which mineral shells are constructed of. Calcium carbonate has two forms, the stable calcite, and the metastable aragonite, which is stable within a reasonable range of chemical environments but rapidly becomes unstable outside this range. When the oceans contain a relatively high proportion of magnesium compared to calcium, aragonite is more stable, but as the magnesium concentration drops, it becomes less stable, hence harder to incorporate into an exoskeleton, as it will tend to dissolve.

With the exception of the molluscs, whose shells often comprise both forms, most lineages use just one form of the mineral. The form used appears to reflect the seawater chemistry – thus which form was more easily precipitated – at the time that the lineage first evolved a calcified skeleton, and does not change thereafter.[7] However, the relative abundance of calcite- and aragonite-using lineages does not reflect subsequent seawater chemistry – the magnesium/calcium ratio of the oceans appears to have a negligible impact on organisms' success, which is instead controlled mainly by how well they recover from mass extinctions.[17] A recently discovered[18] modern gastropod Chrysomallon squamiferum that lives near deep-sea hydrothermal vents illustrates the influence of both ancient and modern local chemical environments: its shell is made of aragonite, which is found in some of the earliest fossil mollusks; but it also has armor plates on the sides of its foot, and these are mineralized with the iron sulfides pyrite and greigite, which had never previously been found in any metazoan but whose ingredients are emitted in large quantities by the vents.[4]

Exoskeleton of a cicada

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Calcium phosphate

Calcium phosphate

The term calcium phosphate refers to a family of materials and minerals containing calcium ions (Ca2+) together with inorganic phosphate anions. Some so-called calcium phosphates contain oxide and hydroxide as well. Calcium phosphates are white solids of nutritious value and are found in many living organisms, e.g., bone mineral and tooth enamel. In milk, it exists in a colloidal form in micelles bound to casein protein with magnesium, zinc, and citrate–collectively referred to as colloidal calcium phosphate (CCP). Various calcium phosphate minerals are used in the production of phosphoric acid and fertilizers. Overuse of certain forms of calcium phosphate can lead to nutrient-containing surface runoff and subsequent adverse effects upon receiving waters such as algal blooms and eutrophication (over-enrichment with nutrients and minerals).

Calcite

Calcite

Calcite is a carbonate mineral and the most stable polymorph of calcium carbonate (CaCO3). It is a very common mineral, particularly as a component of limestone. Calcite defines hardness 3 on the Mohs scale of mineral hardness, based on scratch hardness comparison. Large calcite crystals are used in optical equipment, and limestone composed mostly of calcite has numerous uses.

Aragonite

Aragonite

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

Bryozoa

Bryozoa

Bryozoa are a phylum of simple, aquatic invertebrate animals, nearly all living in sedentary colonies. Typically about 0.5 millimetres long, they have a special feeding structure called a lophophore, a "crown" of tentacles used for filter feeding. Most marine bryozoans live in tropical waters, but a few are found in oceanic trenches and polar waters. The bryozoans are classified as the marine bryozoans (Stenolaemata), freshwater bryozoans (Phylactolaemata), and mostly-marine bryozoans (Gymnolaemata), a few members of which prefer brackish water. 5,869 living species are known. At least two genera are solitary ; the rest are colonial.

Ordovician

Ordovician

The Ordovician is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period 485.4 million years ago (Mya) to the start of the Silurian Period 443.8 Mya.

Polysaccharide

Polysaccharide

Polysaccharides, or polycarbohydrates, are the most abundant carbohydrates found in food. They are long-chain polymeric carbohydrates composed of monosaccharide units bound together by glycosidic linkages. This carbohydrate can react with water (hydrolysis) using amylase enzymes as catalyst, which produces constituent sugars. They range in structure from linear to highly branched. Examples include storage polysaccharides such as starch, glycogen and galactogen and structural polysaccharides such as cellulose and chitin.

Composite material

Composite material

A composite material is a material which is produced from two or more constituent materials. These constituent materials have notably dissimilar chemical or physical properties and are merged to create a material with properties unlike the individual elements. Within the finished structure, the individual elements remain separate and distinct, distinguishing composites from mixtures and solid solutions.

Burrow

Burrow

A burrow is a hole or tunnel excavated into the ground by an animal to construct a space suitable for habitation or temporary refuge, or as a byproduct of locomotion. Burrows provide a form of shelter against predation and exposure to the elements, and can be found in nearly every biome and among various biological interactions. Many animal species are known to form burrows. These species range from small invertebrates, such as the Corophium arenarium, to very large vertebrate species such as the polar bear. Burrows can be constructed into a wide variety of substrates and can range in complexity from a simple tube a few centimeters long to a complex network of interconnecting tunnels and chambers hundreds or thousands of meters in total length; an example of the latter level of complexity, a well-developed burrow, would be a rabbit warren.

Metastability

Metastability

In chemistry and physics, metastability denotes an intermediate energetic state within a dynamical system other than the system's state of least energy. A ball resting in a hollow on a slope is a simple example of metastability. If the ball is only slightly pushed, it will settle back into its hollow, but a stronger push may start the ball rolling down the slope. Bowling pins show similar metastability by either merely wobbling for a moment or tipping over completely. A common example of metastability in science is isomerisation. Higher energy isomers are long lived because they are prevented from rearranging to their preferred ground state by barriers in the potential energy.

Hydrothermal vent

Hydrothermal vent

A hydrothermal vent is a fissure on the seabed from which geothermally heated water discharges. They are commonly found near volcanically active places, areas where tectonic plates are moving apart at mid-ocean ridges, ocean basins, and hotspots. Hydrothermal deposits are rocks and mineral ore deposits formed by the action of hydrothermal vents.

Greigite

Greigite

Greigite is an iron sulfide mineral with the chemical formula Fe2+Fe3+2S4. It is the sulfur equivalent of the iron oxide magnetite (Fe3O4). It was first described in 1964 for an occurrence in San Bernardino County, California, and named after the mineralogist and physical chemist Joseph W. Greig (1895–1977).

Animal

Animal

Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and grow from a hollow sphere of cells, the blastula, during embryonic development. Over 1.5 million living animal species have been described—of which around 1 million are insects—but it has been estimated there are over 7 million animal species in total. Animals range in length from 8.5 micrometres (0.00033 in) to 33.6 metres (110 ft). They have complex interactions with each other and their environments, forming intricate food webs. The scientific study of animals is known as zoology.

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

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Further Reading

Skeleton

Sclerite

Aragonite

Kimberella

Microfossil

Biomineralization

Halkieriid

Evolution of molluscs

Marine invertebrates

Cambrian substrate revolution

Mollusc shell

Nama Group

Shell growth in estuaries

Marine biogenic calcification
See also
References
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External links
Look up exoskeleton in Wiktionary, the free dictionary.
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