Sclerite
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A sclerite (
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
, ', meaning " hard") is a hardened body part. In various branches of biology the term is applied to various structures, but not as a rule to vertebrate anatomical features such as bones and teeth. Instead it refers most commonly to the hardened parts of
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
exoskeleton An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") 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 ...
s and the internal spicules of invertebrates such as certain
sponges Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), 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 t ...
and
soft corals Alcyonacea, or soft corals, are an order of corals. In addition to the fleshy soft corals, the order Alcyonacea now contains all species previously known as "gorgonian corals", that produce a more or less hard skeleton, though quite different ...
. In
paleontology Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
, a scleritome is the complete set of sclerites of an organism, often all that is known from fossil invertebrates.


Sclerites in combination

Sclerites may occur practically isolated in an organism, such as the
sting Sting may refer to: * Stinger or sting, a structure of an animal to inject venom, or the injury produced by a stinger * Irritating hairs or prickles of a stinging plant, or the plant itself Fictional characters and entities * Sting (Middle-earth ...
of a cone shell. Also, they can be more or less scattered, such as tufts of defensive sharp, mineralised bristles as in many marine Polychaetes. Or, they can occur as structured, but unconnected or loosely connected arrays, such as the mineral "teeth" in the
radula The radula (, ; plural radulae or radulas) is an anatomical structure used by molluscs for feeding, sometimes compared to a tongue. It is a minutely toothed, chitinous ribbon, which is typically used for scraping or cutting food before the food ...
of many
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 es ...
, the valves of
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. They are also sometimes known as gumboots or sea cradles or coat-of-mail she ...
s, the beak of
Cephalopod A cephalopod is any member of the molluscan class Cephalopoda ( Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...
, or the articulated exoskeletons of Arthropoda. When sclerites are organised into an unarticulated structure, that structure may be referred to as a scleritome, a term largely used in
paleontology Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
.


Arthropods

In
Arthropoda Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chitin ...
the hardening that produces sclerites is accomplished either by the cross-linking of protein chains in the exocuticle, a process called sclerotization, or by incorporation of minerals such as calcium carbonate into regions of the exoskeleton, or both. Thus, the arthropod exoskeleton is divided into numerous sclerites, joined by less-sclerotized, membranous regions or sutures. Dorsal sclerites of a body segment, often plate-like, are known as '' tergites''. Similarly the ventral sclerites of a body segment are referred to as '' sternites''. Separate sclerites on the lateral aspects of body segments, the
pleura The pulmonary pleurae (''sing.'' pleura) are the two opposing layers of serous membrane overlying the lungs and the inside of the surrounding chest walls. The inner pleura, called the visceral pleura, covers the surface of each lung and dips b ...
, are called ''pleurites''.


Invertebrates other than arthropods

Wide ranges of sclerites of various kinds occur in various invertebrate
phyla Phyla, the plural of ''phylum'', may refer to: * Phylum, a biological taxon between Kingdom and Class * by analogy, in linguistics, a large division of possibly related languages, or a major language family which is not subordinate to another Phy ...
, including
Polychaeta Polychaeta () is a paraphyletic class of generally marine annelid worms, commonly called bristle worms or polychaetes (). Each body segment has a pair of fleshy protrusions called parapodia that bear many bristles, called chaetae, which are ...
and
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 es ...
. Two
taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
that routinely have the term applied however, are the
soft corals Alcyonacea, or soft corals, are an order of corals. In addition to the fleshy soft corals, the order Alcyonacea now contains all species previously known as "gorgonian corals", that produce a more or less hard skeleton, though quite different ...
and the
Porifera Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), 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 t ...
. In both those groups certain of their structures contain mineralised spicules of
silica Silicon dioxide, also known as silica, is an oxide of silicon with the chemical formula , most commonly found in nature as quartz and in various living organisms. In many parts of the world, silica is the major constituent of sand. Silica is ...
or calcium carbonate that are of importance structurally and in defense. Many other invertebrates grow a few hard parts, largely mineralised, as statoliths and similar structures, but those are not generally referred to as sclerites. Clamps, the main attachment structure of the parasitic Polyopisthocotylean
monogenea Monogeneans are a group of ectoparasitic flatworms commonly found on the skin, gills, or fins of fish. They have a direct lifecycle and do not require an intermediate host. Adults are hermaphrodites, meaning they have both male and female repr ...
n flatworms, are composed of various sclerites and associated musculature, located on a posterior organ called the haptor. Clamps are specialized structures attached to the host fish, generally to its gill.


Prehistory

The fossilized scleritome of ''
thumb The thumb is the first digit of the hand, next to the index finger. When a person is standing in the medical anatomical position (where the palm is facing to the front), the thumb is the outermost digit. The Medical Latin English noun for thumb ...
A scleritome is a skeleton made of isolated elements, like the scales of organisms such as the halwaxiids, the teeth in a radula,
spicules Spicules are any of various small needle-like anatomical structures occurring in organisms Spicule may also refer to: *Spicule (sponge), small skeletal elements of sea sponges *Spicule (nematode), reproductive structures found in male nematodes ( ...
in a sponge skeleton, or the elements in a
conodont Conodonts ( Greek ''kōnos'', " cone", + ''odont'', "tooth") are an extinct group of agnathan (jawless) vertebrates resembling eels, classified in the class Conodonta. For many years, they were known only from their tooth-like oral elements, whi ...
apparatus. The term was coined by the palaeontologist Stefan Bengtson. Although sclerites are of considerable importance in the study of extant animals, in palaeontology they are of far greater relative importance because they often are the only parts of an animal that fossilize at all, let alone well or clearly. Many extinct groups are known only from sclerites, leaving moot the question of what their gross anatomy might have looked like. An example of the use of the term in paleontology is to describe hollow calcium carbonate,
calcium sulfate Calcium sulfate (or calcium sulphate) is the inorganic compound with the formula CaSO4 and related hydrates. In the form of γ-anhydrite (the anhydrous form), it is used as a desiccant. One particular hydrate is better known as plaster of Paris ...
or calcium phosphate plates grown as body armor by a number of Early
Cambrian The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ago ...
animals. Unlike
sponge Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), 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 throu ...
spicules, Early Cambrian sclerites appear to be external armor rather than internal structural elements. Sclerites are found on a curious collection of early animals including a common spongelike animal called '' Chancelloria''; an armored slug-like form '' Wiwaxia''; an armored worm with a pair of
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, w ...
-like shells '' Halkieria''; and another armored worm '' Microdictyon'' that is generally considered to be a lobopod/ onychophore. It has been suggested that the sclerites of the Cambrian ''Wiwaxia'' are homologous with the bristles of annelid worms. At least one modern
gastropod The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. T ...
mollusc living near deep sea hydrothermal vents has structures made of iron sulfides similar to some Cambrian sclerites.


References

{{reflist Arthropod anatomy Sponge anatomy Mollusc anatomy