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A scute () or scutum (
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
: ''scutum''; plural: ''scuta'' "
shield A shield is a piece of personal armour held in the hand, which may or may not be strapped to the wrist or forearm. Shields are used to intercept specific attacks, whether from close-ranged weaponry like spears or long ranged projectiles suc ...
") is a bony external plate or scale overlaid with horn, as on the shell of a
turtle Turtles are reptiles of the order (biology), order Testudines, characterized by a special turtle shell, shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Crypt ...
, the skin of
crocodilia Crocodilia () is an order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorph pseudosuchia ...
ns, and the feet of
birds Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
. The term is also used to describe the anterior portion of the
mesothorax The mesothorax is the middle of the three segments of the thorax of hexapods, and bears the second pair of legs. Its principal sclerites (exoskeletal plates) are the mesonotum (dorsal), the mesosternum ( ventral), and the mesopleuron (lateral) ...
in
insect Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...
s as well as some arachnids (e.g., the family Ixodidae, the scale ticks).


Properties

Scutes are similar to
scales Scale or scales may refer to: Mathematics * Scale (descriptive set theory), an object defined on a set of points * Scale (ratio), the ratio of a linear dimension of a model to the corresponding dimension of the original * Scale factor, a number ...
and serve the same function. Unlike the scales of lizards and snakes, which are formed from the
epidermis The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and Subcutaneous tissue, hypodermis. The epidermal layer provides a barrier to infection from environmental pathogens and regulates the ...
, scutes are formed in the lower vascular layer of the skin and the epidermal element is only the top surface . Forming in the living
dermis The dermis or corium is a layer of skin between the epidermis (skin), epidermis (with which it makes up the cutis (anatomy), cutis) and subcutaneous tissues, that primarily consists of dense irregular connective tissue and cushions the body from s ...
, the scutes produce a horny outer layer that is superficially similar to that of scales. Scutes will usually not overlap as
snake scales Snakes, like other reptiles, have skin covered in scales.Boulenger, George A. 1890 The Fauna of British India. p. 1 Snakes are entirely covered with scales or scutes of various shapes and sizes, known as snakeskin as a whole. A scale protects ...
(but see the
pangolin Pangolins, sometimes known as scaly anteaters, are mammals of the order Pholidota (). The one extant family, the Manidae, has three genera: '' Manis'', '' Phataginus'', and '' Smutsia''. ''Manis'' comprises four species found in Asia, while ' ...
). The outer
keratin Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. It is the key structural material making up Scale (anatomy), scales, hair, Nail (anatomy), nails, feathers, horn (anatomy), horns, claws, Hoof, hoove ...
layer is shed piecemeal, and not in one continuous layer of skin as seen in snakes or lizards. The dermal base may contain
bone A bone is a rigid organ that constitutes part of the skeleton in most vertebrate animals. Bones protect the various other organs of the body, produce red and white blood cells, store minerals, provide structure and support for the body, ...
and produce dermal armour. Scutes with a bony base are properly called ''
osteoderm Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, temnospondyls (extinct amph ...
s''. Dermal scutes are also found in the feet of
birds Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
and tails of some
mammals A mammal () is a vertebrate animal of the class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three middle e ...
, and are believed to be the primitive form of dermal armour in reptiles. The term is also used to describe the heavy armour of the
armadillo Armadillos () are New World placental mammals in the order (biology), order Cingulata. They form part of the superorder Xenarthra, along with the anteaters and sloths. 21 extant species of armadillo have been described, some of which are dis ...
and the extinct '' Glyptodon'', and is occasionally used as an alternative to scales in describing
snakes Snakes are elongated Limbless vertebrate, limbless reptiles of the suborder Serpentes (). Cladistically Squamata, squamates, snakes are ectothermic, amniote vertebrates covered in overlapping Scale (zoology), scales much like other members of ...
or certain fishes, such as
sturgeon Sturgeon (from Old English ultimately from Proto-Indo-European language, Proto-Indo-European *''str̥(Hx)yón''-) is the common name for the 27 species of fish belonging to the family Acipenseridae. The earliest sturgeon fossils date to the ...
s, shad, herring, and menhaden.


Mammals

Prehistoric ancestors of mammals, the
synapsids Synapsida is a diverse group of tetrapod vertebrates that includes all mammals and their extinct relatives. It is one of the two major clades of the group Amniota, the other being the more diverse group Sauropsida (which includes all extant rept ...
, are thought to have scutes which were later reduced and replaced by hair. Excluding the attachment surface of fingernails, armoured scutes or scales are almost never seen in modern
mammals A mammal () is a vertebrate animal of the class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three middle e ...
. The horny scales of
pangolin Pangolins, sometimes known as scaly anteaters, are mammals of the order Pholidota (). The one extant family, the Manidae, has three genera: '' Manis'', '' Phataginus'', and '' Smutsia''. ''Manis'' comprises four species found in Asia, while ' ...
s are only rarely called scutes, but "scute" is used to describe the heavy armour of the
armadillo Armadillos () are New World placental mammals in the order (biology), order Cingulata. They form part of the superorder Xenarthra, along with the anteaters and sloths. 21 extant species of armadillo have been described, some of which are dis ...
.


Turtles

The turtle's shell is covered by scutes formed mostly of
keratin Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. It is the key structural material making up Scale (anatomy), scales, hair, Nail (anatomy), nails, feathers, horn (anatomy), horns, claws, Hoof, hoove ...
. They are built similarly to horn, beak, or nail in other species.


Fish

Some fish, such as pineconefish, are completely or partially covered in large armored scales commonly termed scutes. Many Clupeomorpha have an abdominal row of scutes, which are scales with raised, sharp points that are used for protection, and in some cases they also possess dorsal scutes anterior to the dorsal fin, as in '' Knightia'' spp., '' Diplomystus'' spp. and certain extant taxa. Jacks of family Carangidae often have a row of scutes following the
lateral line The lateral line, also called the lateral line organ (LLO), is a system of sensory organs found in fish, used to detect movement, vibration, and pressure gradients in the surrounding water. The sensory ability is achieved via modified epithelia ...
on either side. '' Coelorinchus'' spp. have scutes suborbitally from their preoperculum extending anteriorly onto a triangular snout.
Sturgeon Sturgeon (from Old English ultimately from Proto-Indo-European language, Proto-Indo-European *''str̥(Hx)yón''-) is the common name for the 27 species of fish belonging to the family Acipenseridae. The earliest sturgeon fossils date to the ...
have five rows of scutes—modified ganoid scales—and are otherwise scaleless.


Birds

The
tarsometatarsus The tarsometatarsus is a bone that is only found in the lower leg of birds and some non-avian dinosaurs. It is formed from the fusion of several bird bones found in other types of animals, and homologous to the mammalian tarsus (ankle bones) a ...
and toes of most birds are covered in two types of scales. Large scutes run along the dorsal side of the tarsometatarsus and toes, whereas smaller scutellae run along the sides. Both structures share histochemical homology with reptilian scales; however, work on their evolutionary development has revealed that the scales in bird feet have secondarily evolved via suppression of the feather-building genetic program.Sawyer, R.H., Knapp, L.W. 2003. Avian Skin Development and the Evolutionary Origin of Feathers. J. Exp. Zool (Mol Dev Evol) 298B:57–72. Unblocking the feather suppression program results in feathers growing in place of scales along the tarsometatarsus and toes. Dinosaur species very close to the origin of birds have been shown to have had "hind wings" made of feathers growing from these areas, suggesting that the acquisition of feathers in dinosaurs was a whole-body event. The bottoms of bird feet are covered in small, keeled scale-like structures known as reticulae. Evolutionary developmental studies on these scale-like structures have revealed that they are composed entirely of alpha keratin (true epidermal scales are composed of a mix of alpha and beta keratin). These data have led some researchers to suggest that reticulae are in fact highly truncated feathers.


Insects and other arthropods

The term "scutum" is also used in
insect Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...
anatomy, as an alternative name for the anterior portion of the mesonotum (and, technically, the
metanotum The metathorax is the posterior of the three segments in the thorax of an insect, and bears the third pair of legs. Its principal sclerites ( exoskeletal plates) are the metanotum (dorsal), the metasternum ( ventral), and the metapleuron (late ...
, though rarely applied in that context). In the hard ticks, the Ixodidae, the scutum is a rigid, sclerotised plate on the
anterior Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...
dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage The fus ...
surface, just posterior to the head. In species with eyes, the eyes are on the surface of the scutum. The flexible
exoskeleton An exoskeleton () . is a skeleton that is on the exterior of an animal in the form of hardened integument, which both supports the body's shape and protects the internal organs, in contrast to an internal endoskeleton (e.g. human skeleton, that ...
posterior to the rigid scutum of the female tick, is called the alloscutum, the region that stretches to accommodate the blood with which the mature female tick becomes engorged. Males do not engorge nearly as drastically as females, so they do not need a flexible alloscutum; instead the rigid scutum covers practically the entire dorsal surface posterior to the head, and may be referred to specifically as the conscutum. In some species of
Opiliones The Opiliones (formerly Phalangida) are an Order (biology), order of arachnids, Common name, colloquially known as harvestmen, harvesters, harvest spiders, or daddy longlegs (see below). , over 6,650 species of harvestmen have been discovered w ...
, fused abdominal segments are referred to as a scutum.


See also

* Fish scutes *
Osteoderms Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of Extant taxon, extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, Temnospondyli, ...
*
Scale (zoology) In zoology, a scale (; ) is a small rigid plate (animal anatomy), plate made out of keratin that grows out of Vertebrate animals' skin to provide protection. In lepidopterans (butterflies and moths), scales are plates on the surface of the insec ...
*
Snake scales Snakes, like other reptiles, have skin covered in scales.Boulenger, George A. 1890 The Fauna of British India. p. 1 Snakes are entirely covered with scales or scutes of various shapes and sizes, known as snakeskin as a whole. A scale protects ...
*
Keratin Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. It is the key structural material making up Scale (anatomy), scales, hair, Nail (anatomy), nails, feathers, horn (anatomy), horns, claws, Hoof, hoove ...
*
Skin Skin is the layer of usually soft, flexible outer tissue covering the body of a vertebrate animal, with three main functions: protection, regulation, and sensation. Other animal coverings, such as the arthropod exoskeleton, have different ...
*
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 a rigid outer shell that holds up an organism's shape; the endoskeleton, a rigid internal fra ...


References

{{reflist Animal anatomy Dinosaur anatomy de:Hornschuppe sv:Fjäll (zoologi)